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he  Influence  of  Hemorff 
Unon  Metalioli?;! 


rhiiip  Boi       Hawk. 


The  Influence  of  Hemorrhage 
Upon  Metabolism. 


BY 


Philip  Bovier  Hawk. 


Submitted  in  partial  fulfilment  of  the  requirements  for  the  degree 

of  Doctor  of  Philosophy  in  the  Faculty  of  Pure 

Science,  Columbia  University. 


1905. 

The  Chemical  Publishing  Co., 

Elaston,  Pa. 


a?  17 


CONTENTS. 


PAGE. 

A.  Historical  Introduction. 5 

B.  First  Series  of  Experiments 16 

I.     Description 16 

1 .  The  Purpose  and  Plan  of  the  Investigation 16 

2.  Articles  of  Diet. 17 

3.  Preparation  of  the  Anesthetic 19 

4.  Subject 19 

5.  Methods  of  Analysis 19 

II.     Preliminary  Experiment 20 

III.  First  Hemorrhage 21 

1 .  Operative  Procedure 21 

2.  Observations 23 

3.  Discussion 24 

IV.  Control  Experiments 25 

1.  Influence  of  Anaesthesia 25 

(a).     Observations 26 

( b) .     Discussion 28 

2.  Influence  of  Anaesthesia-operation 29 

(a).     Observations...' 30 

(b).     Discussion 31 

V.     vSecond  Hemorrhage. 32 

1 .  Observations 32 

2.  Discussion 33 

VI.     Third  Hemorrhage 34 

1 .  Observations 35 

2.  Discussion 36 

VII.     Fourth  Hemorrhage 38 

I .     Observations  and  Discussion    39 

VIII.     F'ifth  Hemorrhage 40 

I .     Post  Mortem  Examination 41 

C.  Second  Series  of  Experiments. i 42 

I.     Preliminary  Experiment 42 

II.     Influence  of  Anaesthesia-operation 42 

III.     Influence  of  Hemorrhage. 44 

D.  Alterations  in  the  Specific  Gravity 45 

E.  Relation  Between  Total  Nitrogen  and  Volume  of  the  Urine 46 

F.  Discussion  of  Results. ....    47 

G.  Conclusions 49 

H.     Bibliography 50 

I .     Biographical 53 

J.     Publications 54 


374375 


A.— HISTORICAL  INTRODUCTION. 

The  practice  of  venesection,  or  blood-letting,  is  as  old  almost  as 
tlie  practice  of  medicine.  History  informs  us  that  even  as  early 
as  the  time  of  Galen  the  employment  of  venesection  was  quite 
general.  The  withdrawal  of  blood  was  for  many  centuries 
looked  upon  as  the  most  satisfactory  remedy  possessed  by  the 
practitioner,  and  indeed  in  a  great  number  of  diseases  was  con- 
sidered to  be  the  only  means  of  preserving  life.  The  mere  ques- 
tion of  the  withdrawal  of  blood  was  never  debated,  the  amount 
being  the  only  question  to  decide,  as  it  was  almost  universally 
taken  for  granted  that  some  blood  must  be  drawn.  Galen  himself 
employed  blood-letting  to  the  point  of  syncope,  and  his  principal 
treatment  for  fever  was  blood-letting  and  the  application  of  cold. 
So  great  was  the  faith  in  the  "Universal  treatment,"  as  it  came 
to  be  called,  that  its  domain  of  influence  was  expanded  and  came 
to  include  the  realms  of  religion  and  superstition.  The  desire  to 
do  evil  was  supposedly  dissipated  by  its  magic ;  and,  if  the  vene- 
section were  carried  out  at  the  proper  time,  astrologically,  such  de- 
sirable possessions  as  a  retentive  memory,  a  strong  mind,  etc., 
were  said  to  be  acquired.  The  withdrawal  of  blood  was  accom- 
plished by  means  of  the  leech,  the  cupping  instrument,  or  by  di- 
rect withdrawal  from  an  opened  vein  (venesection),  the  individ- 
ual conditions  as  well  as  the  amount  of  blood  to  be  drawn  gen- 
erally determining  the  metjiod. 

The  universal  practice  of  blood-letting  has,  however,  long  since 
passed,  and  although  there  are  today  certain  diseased  conditions 
in  which  loss  of  blood  is  supposed  to  possess  curative  power,  the 
custom  of  blood-letting  is  practiced  in  much  narrower  limits  than 
in  the  early  centuries  of  its  use.  In  the  words  of  an  eminent  in- 
vestigator "The  waste  of  blood  of  a  not  very  distant  past  is,  let  us 
hope,  gone  forever."  To-day  the  greatest  losses  of  blood  with 
which  we  have  to  deal  occur  either  as  the  result  of  accident,  in 
major  surgical  operations,  or  in  a  few  maladies,  such  as  gastric 
ulcer,  which  are  accompanied  by  internal  bleeding. 

The  present  investigation  was  undertaken  with  the  accidental 
hemorrahge  and  the  loss  of  blood  incident  to  surgical  procedure 


especially  in  mind.  The  variations  in  the  metabolic  activity  as  in- 
dicated by  the  composition  of  the  urine  was  the  principal  point  of 
observation,  and  the  changes  in  the  chemical  composition  of  the 
blood  was  a  secondary  consideration.  No  study  of  the  form  ele- 
ments of  the  blood  was  made. 

It  is  evident  that  a  loss  of  blood  from  any  organism  will  cause 
an  alteration  in  the  physical  and  chemical  composition  of  the  resi- 
dual portion.  It  is  further  well  understood  that  certain  diseases 
are  accompanied  by  changes  in  the  blood.  The  nature  of  those 
alterations  or  changes  (whether  due  directly  to  the  reduction  of 
the  total  blood  or  whether  arising  from  an  internal  pathalogical 
state)  and  their  effect  upon  the  metabolic  functions  of  the  organ- 
ism are  questions  of  great  interest  and  importance.  It  was  but 
natural  that  a  method  of  treatment  holding  undisputed  sway  for 
so  many  hundred  years,  as  did  that  of  blood-letting,  should,  with 
the  growth  of  the  spirit  of  investigation,  be  made  the  subject  of 
many  scientific  observations.  As  would  naturally  follow,  many  of 
these  observations  were  upon  human  subjects,  while  lower  ani- 
mals were  the  subject  of  other  investigations.  Observations  of 
the  first  order  were  obviously  almost  entirely  confined  to  diseased 
subjects.  The  field  covered  by  investigations  upon  the  effect  of 
loss  of  blood  as  well  as  upon  the  influence  of  blood  degeneration 
through  pathalogical  agencies  is  a  field  of  great  breadth,  and 
while  investigations  of  the  second  type  cannot  strictly  be  classed 
with  our  investigation,  yet  for  a  more  complete  survey  of  the  en- 
tire subject  we  have  included  observations  of  that  nature  in  our 
literary  review. 

The  work  of  previous  investigators  may  be  divided  into  three 
general  classes  and  corresponding  sub-groups  : 

Class  I.  Observations  upon  the  influence  of  the  loss  of  blood 
as  shown  by  direct  experimentation  upon  lower  animals. 

1.  Variations  in  the  metabolic  activities  of  the  organism  as 
shown  by  (a)  Examination  of  the  urine  and  determina- 
tion of  the  body  weight. 

(b)  Respiratory  exchange. 

2.  Alterations  in  the  number  of  red  corpuscles  or  percentage 
of  haemoglobin. 

3.  Change  in  the  number  of  leucocytes. 


4-  Influence  on  respiration,  nervous  system,  and  body  tem- 
perature. 
5.  Changes  in  the  fibrin  content,  specific  gravity,  coagulation 
rate,  and  in  tlie  composition  of  the  blood,  serum,  gases,  etc. 
Class  II.  Observations  upon  the  alterations  in  metabolic  activ- 
ity and  in  blood  composition  resulting,  in  human  subjects, 
from  internal  pathalogical  conditions,   (anaemia,  leukaemia, 
chlorosis,  leucocytosis,  etc). 
Class  III.  Observations  upon  the  influence  of  the  loss  of  blood 
occurring  pathalogically  in  human  subjects,    (gastric  ulcer, 
duodenal  ulcer,  post-partum  hemorrhage). 
Our  own  investigation  would  be  classified  under  groups  i   (a) 
and  5  of  the  first  class. 

Investigations  having  for  their  purpose  the  study  of  variations 
in  the  metabolic  activities  of  the  organism  accompanying  hemor- 
rhage as  shown  by  extensive  urine  analyses  are  comparatively  few 
in  number.  The  work  of  Bauer^  is  probably  the  most  frequently 
quoted  in  this  connection.  His  observations  were  upon  well-nour- 
ished and  fasting  dogs,  a  single  experiment  being  made  upon 
each.  A  sheep  hound  weighing  about  20kg.  was  the  subject  of 
the  first  form  of  experiment.  This  animal  was  fed  a  uniform 
daily  diet  until  nitrogen  equilibrium  was  attained,  and  was  then 
subjected  to  a  hemorrhage  of  350-400  cc.  of  blood.  Following 
this  hemorrhage  the  variation  in  the  proteid  metabolism  as  shown 
by  the  nitrogen  content  of  the  urine  was  noted.  The  dog  drank 
water  "ad  libitum."  The  urine  of  the  preliminary  period  showed 
an  average  daily  content  of  16.6  grams  of  nitrogen.  The  hemor- 
rhage caused  an  increased  output  of  nitrogen  and  the  daily  aver- 
age for  the  three  days  following  was  20  grams,  while  the  average 
for  a  period  of  five  days  was  18.9  grams.  Thus  the  immediate 
eflfect  of  a  loss  of  blood  aggregating  about  2%  of  the  body  weight 
of  the  animal  was  an  increase  of  10.2  grams  of  nitrogen  in  three 
days,  and  an  increase  of  11.5  grams  in  five  days.  From  this  point 
the  rate  of  excretion  of  nitrogen  fell  rapidly,  being  slightly  below 
normal  at  the  end  of  the  experiment.  The  volume  of  the  urine 
increased  somewhat  after  blood-letting,  probably  due  to  uncon- 
trolled water  ingestion,  while  the  animal  also  gained  slightly  in 
body  weight. 


8 

Experiments  on  a  fasting  dog  were  also  followed  by  a  rise  in 
the  urine  volume  and  in  the  nitrogen  content.  From  the  data  the 
author  concluded  that  the  influence  of  the  loss  of  blood  was  much 
greater  in  the  well-nourished  organism  than  in  the  organism  in 
the  fasting  condition. 

As  a  "check"  upon  his  results  Bauer  made  three  control  experi- 
ments in  which  the  customary  operative  steps  were  followed  but 
no  blood  was  drawn.  These  were  on  well-nourished  dogs  in  a 
condition  of  nitrogen  equilibrium  and  from  them  he  concluded 
that  the  operation  had  no  influence  upon  the  execution  of  nitro- 
gen by  the  urine. 

Experiments  along  lines  similar  to  those  pursued  by  Bauer  have 
been  made  by  Jiirgensen,-  Maltschewsky,^  and  Ascoli  and 
Draghi.*  The  general  conclusions  of  Jiirgensen  are  the  same  as 
those  of  Bauer.  He  states  that  the  increase  in  nitrogen  in  the  case 
of  the  well-nourished  animals  is  absolutely  greater  after  loss  of 
blood  than  in  fasting  animals,  but  that  it  is  smaller  in  comparison 
with  the  quantity  of  proteid  available.  Jiirgensen's  experiments 
are  interesting  but  his  failure  to  give  his  data  for  the  control  ex- 
periments upon  dogs  in  a  condition  of  nitrogen  equilibrium  is  to 
be  regretted. 

Fraenkel'^  takes  exception  to  the  conclusions  of  Bauer  and  Jiir- 
gensen, and  claims  that  the  increased  proteid  decomposition  fol- 
lows the  diminished  oxygen  supply,  due  to  the  lowering  of  the 
oxygen  carrying  power  of  the  blood.  This  theory  is  not  approved 
by  v.  Noorden,**  Jiirgensen  and  others. 

Ascoli  and  Draghi,  in  observations  upon  bmman  subjects  and 
upon  dogs,  came  to  the  conclusion  that  there  was  no  increased 
nitrogen  elimination  following  the  loss  of  blood.  The  non-occur- 
ence  of  a  rise  in  nitrogen  after  hemorrhage  is  based  on  the  theory 
that  the  organism  contains  a  certain  reserve  supply  of  blood  which 
may  be  removed  without  detriment  to  the  body  (the  "luxus  blood" 
of  Maragliano). 

Observations  upon  the  variation  in  the  respiratory  exchange, 
after  hemorrhage,  have  been  made  by  Bauer,  Lukjanow,'^  and 
Giirber.^  Experiments  by  Bauer  in  a  respiration  apparatus  of  the 
Voit  model,  using  fasting  dogs  as  subjects,  showed  the  excretion 
of  COo  to  be  practically  unchanged  in  the   12  hours  following 


hemorrhage,  while  the  oxygen  absorption  decreased  about  15%. 
Forty-eight  hours  later  the  amounts  of  both  CO-  excreted  and  O 
absorbed  had  decreased  greatly.  In  well-nourished  dogs  there 
was  an  increase  of  4%  in  CO^  excreted  during  the  first  12  hours 
after  hemorrhage  and  an  increase  of  22^0  in  the  oxygen  absorbed. 
After  24  hours  the  values  for  both  CO^  and  O  sank  appreciably. 
Bauer  concluded  that  the  decrease  in  CO^  excretion  after  hemor- 
rhage showed  that  the  destruction  of  fat  had  decreased  and  point- 
ed to  the  storing  of  fat  in  anaemic  and  chlorotic  patients  as  veri- 
fication. 

GiJrber,  as  a  result  of  observations  on  well-nourished  rabbits, 
drew  the  conclusion  that  the  respiratory  exchange  w^as  eiit'ected 
very  little  if  any  after  hemorrhage,  and  that  the  variation  when 
observed  was  in  the  form  of  an  increase.  Lukjanovv'  investigated 
the  eftect  of  oxygen  inhalation  after  hemorrhage.  Two  experi- 
ments were  made  on  rats  and  one  on  a  dog.  In  every  case  one 
hour  after  the  hemorrhage  the  absorption  of  oxygen  was  in- 
creased. 

Among  the  numerous  observations  upon  the  alteration  in  the 
number  of  red  corpuscles  or  the  percentage  of  haemoglobin  fol- 
lowing hemorrhage  those  of  Autokonenko,''  Baumann,"'  Luzet,^^ 
X'ulpain.'-  Hiinerfauth.^"  Buntzen'*  and  v.  Lesser"  are  of  par- 
ticular interest.  All  of  the  experiments  of  Autokonenko  were 
made  on  dogs.  His  animals  w-ere  always  fed  a  constant  diet  be- 
tween 9  and  TO  A.  M.  and  the  operations  occurred  between  3  and 
4  P.  M.  The  author  concluded  that  the  number  of  leucocytes  be- 
gan to  increase  from  the  first  hour  after  blood-letting,  the  maxi- 
mum being  reached  at  the  end  of  the  first  day.  The  increase  w^as 
principally  in  small  leucocytes.  If  a  second  hemorrhage  was  in- 
duced when  the  leucocytes  were  still  greatly  increased  a  leucocy- 
tosis  more  prominent  than  the  first,  but  of  less  duration,  occurred. 
A  third  hemorrhage  produced  no  further  increase.  Baumann  in 
a  series  of  very  complete  experiments  on  dogs  observed  a  general 
deterioration  in  the  blood  and  serum  following  hemorrhage,  es- 
pecially in  the  former.  The  quantity  of  haemoglobin  he  found  to 
be  reduced  21%,  the  number  of  red  corpuscles  14%,  and  the  spe- 
cific gravity  1%,  The  solids,  proteids  and  total  nitrogen  of  the 
blood  and  serum  underwent  a  decrease  of  about  12%  each.     The 


lO 

leucocytes  were  increased  about  40%.  Serum  albumin  was  found 
to  have  been  greatly  increased  at  the  expense  of  the  serum  globu- 
lin. Fibrin  was  increased  and  the  coagulation  rate  shortened 
while  the  ash  remained  unchanged. 

Luzet,  as  the  result  of  investigations  on  pigeons,  found  a  de- 
crease in  red  corpuscles  and  haemoglobin  and  an  accompanying 
leucocytosis.  The  work  is  of  interest  in  indicating  the  similarity 
of  post-hemorrhagic  effects  in  man,  lower  animals  and  birds.  The 
conclusions  of  Buntzen  from  experiments  on  dogs  are  interest- 
ing:— I.  Red  corpuscles  decreased  5%  in  10  minutes.  2.  Dia- 
meter of  red  corpuscles  decreased.  3.  No  increase  in  leucocytes. 
4.  Volume  of  blood  restored  in  a  few  hours  after  a  hemorrhage 
of  1-2%  body  weight.  Loss  of  over  4%  took  over  24  hrs.  to  re- 
store. 5.  Red  corpuscles  normal,  after  hemorrhage  of  1.1%  to- 
4.4%  of  body  weight,  in  7  to  34  days.  6.  Percentage  of  solids  in 
the  serum  decreased  from  11.5%  to  10.1%  in  successive  draw- 
ings. 

The  failure  of  the  author  to  observe  any  increase  in  leucocytes 
after  hemorrhage  is  contradictory  to  nearly  every  authentic  obser- 
vation. 

Hiinerfauth  investigated  the  influence  of  "traumatic  anaemia" 
in  frogs,  rabbits  and  dogs.  The  observations  on  frogs  show  that 
in  an  hour  after  a  hemorrhage  of  less  than  2%  body  weight  a 
very  noticeable  decrease  in  the  number  of  red  cells  was  seen,  and 
a  lesser  decrease  in  amount  of  haemoglobin.  A  strong  leuco- 
cytosis also  existed.  Regeneration  was  shown  to  be  slow  in  the 
frog.  Regeneration  in  the  frog  was  also  studied  by  Vulpain,  who 
found  it  to  be  much  slower  than  in  man  and  the  lower  animals. 
Six  weeks  after  the  hemorrhage  the  cells  in  the  marrow  destined 
to  form  new  red  corpuscles  were  small  and  totally  colorless. 

Investigations  of  more  or  less  value  have  been  carried  out  by 
Lyon,^''  Bizzozero  and  Salvioli,^^  Monassein,^^  Tschudnowsky,^^ 
Rieder,^"  and  Hayem.-^  Bizzozero  and  Salvipli  claim  that  a  hem- 
orrhage of  1%  body  weight  produced  a  decrease  of  11.14%  ii^  the 
haemoglobin  and  that  this  ratio  holds  generally.  In  a  series  of 
observations  on  a  large  number  of  different  subjects  (leech, 
mouse,  mole,  rabbit,  hare,  hen  and  pigeon)  Monassein  found  the 


II 

red  corpuscles  to  increase  in  size  after  hemorrhage.  He  explain- 
ed this  as  being  due  to  an  imbibition  of  the  diluted  plasma. 

Investigations  made  upon  animals,  in  which  the  change  in 
the  number  of  leucocytes  following  hemorrhage  was  ob- 
served, have  been  promoted  by  Hayem,  Rieder,  Himmelj- 
sterna."--  Tschoudnowsky,  Virchow,-^  Limbeck,-*  and  Molas- 
sez.-"'  Hayem  observed  a  leucocytosis  which  he  ascribes  as 
arising  rather  from  the  inflammatory  process  than  from 
the  loss  of  blood.  Limbeck  observed  a  similar  leuco- 
cytosis. and  also  found  an  increase  in  leucocytes  following 
simple  incision,  when  no  blood  was  drawn  and  the  wound  healed 
with  no  suppuration.  Tschoudnowsky  found  such  a  rapid  leuco- 
cytosis that  the  ratio  of  leutocytes  to  red  corpuscles  was  as  high 
as  1 :6o.  \"irchow  explained  post-hemorrhagic  leucocytosis  as  be- 
ing caused  by  the  property  the  leucocytes  have  of  clinging  to  the 
walls  of  the  vessels  when  the  other  constituents  are  drawn  oft.  In 
a  series  of  experiments  on  dogs  Rieder.observed  a  variable  leuco- 
cytosis after  hemorrhage.  In  one  case  the  number  of  leucocytes 
was  double  the  normal  on  the  third  day,  while  in  another  case  it 
was  very  feeble  at  that  period.  Himmelj  sterna  reported  the 
unique  observation  of  a  decrease  in  leucocytes  on  the  first  day 
after  hemorrhage. 

Henle,-''  Remak.-'  W'oltersom,-*  Conheim,'^  Massart  and  Bor- 
det,-'-'  Gabritschewsky,-''^  Afanasiew,^-  Leber,-''^  Roemer,^*  Buch- 
ner,-"-''  Escherich,-'*'  and  Foster  "'  all  report  the  observation  of  a 
leucocytosis  of  greater  or  lesser  intensity  following  hemorrhage. 
Afanasiew  reports  an  increase  in  the  leucocytes  amounting  to  15 
times  the  normal. 

The  influence  of  the  loss  of  blood  upon  the  nervous  system  has 
been  investigated  by  Leichtenstern,^^  Kussmaul  and  Tenner, ^^ 
Naunyn  and  Quinke,*"  and  Gies."*^  Leichtenstern  observed  that 
immediately  after  hemorrhage  a  temporary  decrease  in  the  num- 
ber and  depth  of  the  respirations  took  place.  After  severe  hemor- 
rhages the  frer|uency  of  respiration  ultimately  increased.  Kuss- 
maul and  Tenner  observed  convulsions  due  to  a  decrease  in  the 
aterial  blr)od  supply  of  the  brain.  These  convulsions  did  not  occur 
in  slow  hemorrhages.  The  authors  showed  the  central  origin  of 
the  convuslions  to  be  in  the  region  l)ehind  the  optic  lobes.     This 


12 

investigation  of  Kussmaul  and  Tenner,  as  well  as  others  by  Jolly  ' 
and  Bonders,  has  shown  that  the  statement  that  the  loss  of  blood 
does  not  alter  the  blood  supply  to  the  brain  is  without  foundation. 
In  experiments  on  toads,  frogs,  rabbits  and  dogs  Gies  induced 
anaemia  of  the  brain  by  means  of  perfusing  solutions.  He  ob- 
served that  rapidly  induced  anaemia  was  followed  by  convulsions, 
whereas  an  anaemia  brought  about  gradually  was  unaccompanied 
by  any  such  phenomena. 

Experiments  upon  the  variations  in  blood  pressure  after  hem- 
orrhage have  been  made  by  Worm-MuUer,*-  Volkmann,*''  Naw- 
rotzky,"*  Magendie,*'^  Nawlichen**^  and  Hayem,*'  while  observa- 
tions upon  the  variations  in  body  temperature  after  hemorrhage 
have  been  made  by  Marshall  Hall,**  Traube,*^  Spielman,-''*'  Frese 
and  Charaszewski,^^  and  Wunderlich.^- 

The  fibrin  content  of  the  blood  has  been  made  the  subject  of 
investigations  by  Biicke,^'^  Nasse,''*  Jiirgensen,^^  and  Mayer.^** 
Briicke  found  a  decrease  in  fibrin  proportional  to  the  amount  of 
depletion.  His  observations  were  made  upon  several  samples, 
drawn  consecutively.  He  attributed  the  decrease  in  fibrin  to  the 
fact  that  transuding  fluids  had  diluted  the  blood.  Nasse,  by  draw- 
ing the  blood  at  intervals  of  48  hours,  showed  an  increase  in  fibrin, 
and  Jiirgensen  by  similar  procedure  on  fasting  dogs,  confirmed 
this  observation.     Baumann  and  Mayer  found  the  same. 

The  rate  of  coagulation  has  been  studied  by  Vierordt," 
Briicke,'^^  Nasse,^^  and  Baumann.*"'  Vierordt  and  Briicke 
reached  the  conclusion  that  in  uninterrupted  hemorrhages  the 
coagulation  rate  gradually  shortened.  Nasse,  by  drawing  his 
samples  at  intervals  of  24  hovirs,  found  the  coagulation  rate  to 
lengthen.  Baumann  after  an  interval  of  a  week  found  the  coag- 
vilation  rate  shortened.  Probably  much  depends  upon  the  individ- 
ual organism  and  the  amount  of  blood  drawn. 

Observations  upon  the  specific  gravity  of  the  blood  have  been 
made  by  Sherrington  and  Copeman,"^  Otto,"-  Jones,*'-^  Bizzozero 
and  Salvioli,"*  Woltersom,*''*  and  Baumann.''"  In  every  case  the 
specific  gravity  was  observed  to  fall  after  loss  of  blood. 

Variations  in  body  weight  after  hemorrhage  have  been  noted 
by  TolmatschefT,"^  Bauer,"'*  Jiirgensen,"''  Lister,'^"  and  Hiiner- 
fauth.'^^     An  increase  was  generally  observed.     Tolmatschefif  in- 


13 

duced  six  hemorrhages  upon  one  dog  in  a  period  of  70  days  and 
observed  an  increase  in  body  weight  up  to  the  fifth  hemorrhage, 
when  a  decrease  occurred.  His  hemorrhages  were  from  1.1%  to 
2.8%  of  the  body  weight.  In  experiments  already  quoted  upon 
nitrogen  metabolism  Bauer  and  Jiirgensen  observed  a  slight  rise 
in  body  weight.  Lister  mentions  the  fattening  of  calves  in  Eng- 
land through  frequent  blood-letting.  Hiinerfauth  observed  in 
dogs  an  increase  in  weight  during  the  week  after  hemorrhage. 

The  rate  of  the  urine  flow  after  hemorrhage  has  been  investi- 
gated by  Frederick  Goll.'-  Dogs  narcotized  by  opium  were  used 
as  subjects.  Ureter  cannulas  were  inserted  and  the  normal  flow 
of  urine  was  observed  for  one-half  hour  when  a  hemorrhage  was 
induced  and  the  variations  in  the  flow  of  urine  and  in  blood  pres- 
sure were  recorded.  In  every  case  the  urine  flow  was  greatly  de- 
creased after  hemorrhage,  and  this  decrease  was  accompanied  by 
a  fall  in  blood  pressure.  The  author  concluded  from  his  obser- 
vations that  arterial  pressure  had  an  important  influence  upon  the 
excretion  of  urine.  Jiirgensen  observed  an  increased  urine  flow 
after  hemorrhage,  especially  in  the  case  of  fasting  dogs.  He  of- 
fered two  suggestions  for  the  increased  volume: — i.  Extra  water 
needed  to  remove  the  accumulated  urea.  2.  Tissue  fluids  contain 
less  proteid  and  more  water  than  the  blood.  By  transudation 
after  hemorrhage  the  blood  would  contain  more  water  than  nor- 
mal if  it  were  not  removed  in  the  urine. 

"Fatal  hemorrhage"  has  been  the  topic  of  investigations  by 
Schramm,  v.  Kireefif,'^  Hayem,'^  and  Noll.'"  Schramm  placed 
the  limit  as  varying  between  3.91% — 5-77%  of  the  body  weight, 
while  Hayem  placed  the  value  at  from  4.34% — 5-55%  depending 
on  the  individual,  v.  Kireeff  ascertained  that  a  loss  of  between 
56%  and  95%  of  the  total  blood  was  necessary  to  cause  death, 
whereas  Xoll  foimd  that  a  hemorrhage  of  -A,  the  total  blood  was 
fatal. 

Infusion  experiments  of  various  forms  have  been  promoted  by 
Kronecker,"  Ott.''*  Maydl,''*  Schramm,'"'  and  Prevost  and 
Dumas. ^' 

The  influence  of  iron  feeding  u])on  blood  regeneration  follow- 
ing hemorrhage  has  been  investigated  by  Skvortsov,"^  and  De- 
bierre  and  Ij'nassier.'*''     The  results  of  both  observations  showed 


14 

an  increase  in  haemglobin  and  red  corpuscles  after  iron  feeding 
was  begun.  An  increase  in  lymph  formation  was  obtained  by 
Emminghaus^*  after  hemorrhage. 

Proteid  decomposition  in  patients  suffering  from  various  mala- 
dies has  been  studied  by  Fleischer  and  Penzoldt,^^  Lipmann- 
Wulf,**'  Ketcher,*'  v.  Moraczewski,***  and  Eichorst.*'  The  obser- 
vations of  Fleischer  and  Penzoldt  were  concerned  with  conditions 
pertaining  in  Leukaemia  patients.  Normal  persons  were  used  as 
controls  and  were  fed  the  same  diet  as  the  patients.  The  urine 
was  analyzed  for  nitrogen,  sulphur,  phosphorus  and  uric  acid. 
The  data  showed  that  all  four  of  these  constituents  were  increased 
in  leukaemia.  Lipmann-Wulf  in  three  cases  of  chlorosis  observed 
a  gain  of  nitrogen  to  the  body  varying  from  0.06  to  0.71  gram 
per  day.  However,  the  nitrogen  content  of  the  food  was  deter- 
mined by  calculation,  and  the  feces  was  not  analyzed.  Eichorst, 
on  the  contrary,  claims  to  have  observed  an  increased  proteid  de- 
composition in  pernicious  anaemia.  He  simply  makes  his  deduct- 
ions from  general  indications  as  no  food  analyses  were  made. 

The  effect  of  the  inhalation  of  oxygen  in  diseases  where  a  path- 
alogical  blood  condition  maintains  has  been  investigated  by  Bur- 
zhinski,^°  Sticker,^^  Albrecht,^-  and  Hayem.^^  The  observations 
of  Burzhinski  show  an  increased  nitrogen  metabolism  in  leukae- 
mia patients.  Uric  acid  in  relation  to  urea  was  also  increased. 
Studies  by  Aibrecht  on  anaemic  children  who  inhaled  thirty  litres 
of  oxygen  daily  showed  increased  respiration,  pulse,  body  tem- 
perature and  weight.  The  red  corpuscles  were  increased  in  di- 
rect ratio  to  the  amount  of  oxygen  inhaled  and  the  haemoglobin 
value  was  also  raised. 

Post-hemorrhagic  observations  of  leucocytes  have  been  made 
by  Rieder,''*  Hayem,'^''  Samuel,'"'  and  King.^^  Rieder  observed 
that  the  leucocytosis  following  hemorrhage  in  man  did  not  differ 
from  other  forms  of  leucocytosis.  He  found  nucleated  red  cor- 
puscles and  a  decrease  in  haemoglobin.  Hayem  failed  to  observe 
any  post-hemorrhagic  leucocytosis  in  human  subjects.  The  red 
corpuscles  and  haemoglobin  were  decreased.  Samuel  observed 
that  the  number  of  the  leucocytes  was  increased  after  hemor- 
rhage, due  to  the  large  number  which  were  brought  by  the  lymph. 

Post-operative  leucocytosis  was  observed  in  several  instances 


i 


by  King-.  The  operations  were  of  various  kinds  of  the  major 
order.  The  maximum  leticocytosis  in  the  majority  of  cases  oc- 
curred within  twelve  hours  after  the  operation  and  was  very 
transient.  The  author's  observation  of  leucocytosis  following 
operative  procedure  agrees  with  those  of  ]\Iolassez,  and  Hayem. 
These  latter  investigators  however  were  inclined  to  believe  the 
leucocytosis  was  due  to  suppuration  or  inflammation,  whereas  no 
such  condition  was  present  in  King's  experiments. 

Mosler,'-'*  Lowit,^''  and  Miiller^""  have  observed  conditions  in 
leukaemia.  Low  it  claimed  that  the  large  nimiber  of  leucocytes  in 
the  blood  in  leukaemia  was  not  due  to  increased  production  of  leu- 
cocytes but  to  a  lessening  of  the  formation  of  polynuclear  cells 
from  the  mononuclear,  as  well  as  to  the  diminution  in  the  destruc- 
tion of  pol\nuclear  cells.  Miiller  believed  the  increase  in  leuco- 
cytes was  due  to  increased  cell  proliferation. 

The  observations  upon  the  influence  of  the  loss  of  blood  occur- 
ring pathalogically  in  human  subjects  are  comparatively  few  in 
number.  Such  losses  would  ordinarily  occur  from  accident,  in 
rupture  of  aneurism,  in  post-partum  hemorrhage,  or  in  cases  of 
ulcer  of  the  stomach  or  duodenum.  Osler"^  mentions  an  instance 
where  /JX  pounds  (3375  grams)  of  blood  was  shed  into  the 
pleura  from  the  rupture  of  an  aneurism.  In  a  case  of  hemateme- 
sis  the  same  authority  mentions  that  a  loss  of  10  pounds  (4500 
grams)  in  one  week  was  followed  by  recovery.  The  author  states 
that  even  after  very  severe  hemorrhages  of  this  order,  the  num- 
ber of  red  corpuscles  is  not  reduced  so  greatly  as  in  forms  of 
idiopathic  of  pernicious  anaemia.  Thus  in  the  above  instance 
after  a  week  of  bleeding  the  red  corpuscle  count  was  as  high  as 
1,390,000  per  cubic  millimetre.  Regeneration  goes  on  rapidly  and 
sometimes  the  blood  is  normal  as  regards  volume  and  content  of 
salts  and  proteid  constituents,  in  a  week  or  ten  days.  This  regen- 
eration may,  however,  take  weeks  or  even  months,  Osier  says,  be- 
fore the  corpuscles  reach  the  normal  standard.  The  haemoglobin 
is  restored  more  slowly  than  the  corpuscles,  and  there  is  a  mod- 
erate leucocytosis  which  diminishes  during  recovery. 

Nitrogen  metabolism  during  gastric  ulcer  was  studied  by  v. 
Noorden,'"^  Xeusser,'"'*  and  Kalisch.'"^  v.  Noorden  observed  in 
two  cases,  where  the  loss  of  blood  was  large,  that  no  considerable 


i6 

increase  was  noted  in  the  nitrogen  elimination,  either  on  the  day 
of  the  loss  of  blood  or  on  the  following  days.  The  patients  took 
no  food  and  the  nitrogen  output  was  from  6.2  to  8  grams  per  day, 
an  amount  which  was  about  normal  for  the  organisms  during 
inanition.  From  this  the  author  was  inclined  to  doubt  whether 
the  increase  in  nitrogen  observed  by  Bauer,  Jiirgensen  and  others 
w^ould  hold  in  the  case  of  man. 


B.— FIRST  SERIES  OF  EXPERIMENTS. 


I.     DESCRIPTION. 

I.  The  Purpose  and  Plan  of  the  Investigation. — The  purpose  of 
this  investigation  was  to  study,  in  dogs,  the  effect  of  external 
hemorrhage  upon  the  general  metabolic  processes  of  the  body, 
particular  attention  being  paid  to  the  changes  occurring  in  the 
volume  of  the  urine  and  in  the  course  of  the  nitrogen,  sulphur  and 
phosphorus  output.  It  was  also  proposed  to  study  the  variation  in 
the  chemical  composition  of  the  blood  following  hemorrhage. 
Alterations  in  body  weight  were  also  recorded.  It  was  deter- 
mined to  use  one  subject  for  a  series  of  experiments  and  to  study 
the  influence  of  repeated  hemorrhages  extending  through  a  long 
period  of  time,  and  later  to  check  these  results  upon  a  second 
animal. 

After  the  first  dog  was  brought  to  a  condition  of  nitrogen  equi- 
librium by  means  of  a  preliminary  experiment,  the  initial  hemor- 
rhage was  instituted.  Following  this,  after  the  dog.  was  again  at 
nitrogen  equilibrium  came  the  control  experiments.  These  were 
followed  by  four  more  hemorrhages  of  varying  intensities  and 
separated  from  one  another  by  periods  of  different  lengths.  The 
first,  second,  third  and  fourth  hemorrhages  were  made  with  the 
animal  practically  in  nitrogen  equilibrium,  w^hereas  in  the  case  of 
the  fifth  hemorrhage  no  attempt  in  this  direction  was  made.  We 
fully  appreciate  that  at  the  time  of  this  last  hemorrhage  the  or- 
ganism was  somewhat  abnormal  and  that  therefore  the  results  are 


I? 

not  strictly  comparable  with  those  of  the  earlier  hemorrhages. 
However,  as  long  as  we  were  able  to  get  the  organism  in  a  condi- 
tion of  nitrogen  equilibrium  we  hold  that  no  great  abnormality 
could  obtain,  and  that  therefore  we  are  justified  in  comparing  the 
data  from  the  first  four  hemorrhages. 

General  data  for  the  first  series  of  experiments  are  given  in 
Table  XII. 

2.  Articles  of  Diet.  Beef. — This  consisted  of  lean  meat  (round 
steak )  carefully  freed,  as  far  as  possible  mechanically,  from  all 
traces  of  fat  and  connective  tissue.  After  being  finely  hashed  in 
a  meat  chopper  the  meat  was  thoroughly  pressed  in  a  hand  press, 
carefully  mixed  and  samples  taken  for  analysis.  This  sampling 
was  done  by  taking  small  portions  from  various  parts  of  the  mass 
during  the  mixing  process  and  weighing  by  difference  in  tubes 
previously  freed  from  moisture  by  means  of  filter  paper.  After 
being  pressed  and  mixed  the  beef  was  made  into  small  balls  and 
placed  in  air-tight  glass  jars.  These  vessels  contained,  at  most, 
only  enough  nieat  for  two  days'  use,  thus  necessitating  the  open- 
ing of  any  single  jar  but  once. 

The  meat  was  prepared  in  quantity  sufficient  to  last  several 
weeks,  and  for  preservation  was  placed  in  the  "cold  room"  where 
it  was  frozen,  thus  insuring  uniform  composition  from  day  to  day. 
Proof  of  this  uniform  composition  after  the  lapse  of  time  was 
shown  by  the  duplication  of  analytic  results  three  weeks  after  the 
beef  had  been  placed  in  the  jars.  We  feel  that  this  method  serves 
admirably  for  the  purpose  intended.  The  beef  is  at  all  times  fresh 
and  palatable  and  eagerly  eaten  by  the  animals  under  investiga- 
tion. In  any  experunents  where  use  is  made  of  fresh  meat  and 
where  freezing  facilities  are  at  the  command  of  the  investigator, 
we  highly  recommend  this  method  of  meat  preservation  and 
preparation.' 

Cracker  Dust. — Ordinary  cracker  dust  purchased  in  quantity 
from  a  wholesale  grocer,  thoroughly  mixed  and  placed  in  large 
air-tight  anatomical  jars. 

Lard. — Pure  leaf  lard  and  containing  nothing  of  a  vegetable 
nature. 

Bone  Ash. — Obtained  from  a  well    known  firm  of    wholesale 

ICies  :     American  Journal  of  Physiology,  1501,  V.,  p.  235. 


i8 

chemists.  The  feces  of  dogs  hving  upon  a  meat  diet  are  ordinar- 
ily passed  in  large  quantity  at  intervals  of  several  days  and  being 
of  a  very  soft  character  are  extremely  unpleasant  to  deal  with. 
Feces  of  this  order  will  invariably  become  distributed  over  a 
great  portion  of  the  interior  of  the  cage  and  frequently  mix  with 
the  urine,  thus  making  impossible  accurate  deductions  regarding 
the  nitrogen  content  of  either  the  one  or  the  other.  In  order  to 
cause  more  regular  defecation  of  feces  'having  a  more  desirable 
consistenc}'  it  was  determined  to  feed  the  animal  ten  grams  of 
bone  ash  daily.  With  this  amount  of  ash  in  the  diet  the  animal 
defecated  eighty  times  in  eighty-four  days  or  approximately  once 
a  day,  and  with  two  exceptions  the  stools  were  invariably  of  a 
hard  character  and  easily  removable. 

Water. — Ordinary  city  water  heated  to  about  fifty  degrees  cen- 
tigrade. The  water  was  taken  at  this  high  temperature  in  order 
to  offset  the  low  temperature  of  the  frozen  beef  and  at  the  same 
time  form  a  tempting"  mixture.  When  added  to  the  solid  ingre- 
dients of  the  diet  this  warm  water  formed  a  rather  thick  soup 
which  was  decidedly  appetizing.  The  food  mixture  as  finally 
prepared  possessed  a  temperature  of  approximately  twenty-three 
degrees  centigrade  and  was  always  eaten  by  the  dog  with  great 
eagerness. 

By  referring  to  Table  I,  the  quantity  of  each  of  the  articles  of 
diet  fed  during  the  various  periods  may  be  observed.  This  diet 
containing  10.271  grams  of  nitrogen,  was  given  the  beast  at  5 
o'clock  every  afternoon.  It  was  of  course  very  desirable,  from 
the  nature  of  the  investigation,  that  the  subject  should  ingest  the 
same  amount  of  nitrogen  daily  the  entire  experimental  period. 
Therefore,  when  the  original  preparation  of  beef  was  consumed, 
enough  of  the  new  preparation  was  fed  to  give  the  same  amount 
of  nitrogen  daily.  Thus  the  different  beef  preparations  were  fed 
as  follows : — 

Preparation  No.  i. — 250  grams  daily  from  Nov.  2  to  Dec.  4. 

Preparation  No.  2. — 256.4  grams  daily  from  Dec.  5  to  Jan.  2. 

Preparation  No.  3. — 241  grams  daily  from  Jan.  3  to  Jan.  24. 

Apart  from  the  beef  the  amounts  of  the  constituents  of  the 
diet  fed  daily  were  unaltered  during  the  entire  experimental 
period  of  eighty-four  days. 


19 

3-  Preparation  of  the  Anaesthetic. — The  ether  used  in  all  oper- 
ations was  the  purest  product  obtainable  on  the  market.  This 
presumably  chemically  pure  ether  was  dehydrated  by  fused  copper 
sulphate  for  several  days,  and  after  being  subjected  to  a  series  of 
three  distillations  the  final  product  was  that  used  as  the  anaes- 
thetic in  our  operations. 

The  chloroform  was  also  the  best  obtainable  and  was  used  as 
purchased. 

4.  Subject. — In  all  long  metabolism  experiments  upon  dogs 
where  so  much  of  the  success  of  the  investigation  depends  upon 
the  subject  it  is  evidently  of  great  importance  to  secure  a  dog 
that  will  eat  his  food  regularly,  be  contented  in  confinement,  and 
possess  an  amiable  disposition.  Where  the  income  and  outgo  of 
nitrogen  are  studied  it  is  also  desirable  to  use  a  short-haired  dog, 
as  in  the  course  of  a  long  experiment  much  nitrogen  is  lost  by 
way  of  the  hair.  In  the  present  instance  three  beasts  were  tried 
before  a  suitable  subject  was  found.  The  third  animal,  however, 
a  very  active  short-haired  dog  weighing  about  17k.  was  entierly 
satisfactory. 

5.  Methods  of  Analysis. — The  nitrogen  determinations  were 
made  by  the  Kjeldahl  method,  the  preliminary  oxidation  being  ac- 
coir.plished  by  sulphuric  acid  and  a  small  amount  of  copper  sul- 
phate.^ The  chlorine  content  of  the  urine  was  determined  by 
Mohr's-  method.  Sulphur  and  phosphorus  were  determined  by 
the  fusion  method."'  The  specific  gravity  of  the  blood  and  urine 
was  determined  by  an  urinometer. 

The  analyses  in  every  instaiKe  were  made  in  duplicate. 

The  analytic  results  were  further  controlled  by  the  preparation 
and  analysis  of  composite  urine  samples  for  each  experimental 
period.  In  Table  XIY  are  given  the  data  from  the  analyses  of 
these  urines. 

Xothing  except  the  [)urest  chemicals  available  were  employed 
and  these  were  always  examined  and  "checked"  before  use. 

1  Marcuse  :    Archiv  fiir  die  gesammte  Physiologic,  1896.  LXIV.,  p.  2.^2. 

2  Neubauer  and  Vogel :    Harnanalyse  (Modification  of  Neubauer  and  Salkowski),  1S98, 
p.  709- 

'Hawk  :     University  of  Pennsylvania  Medical  Bulletin,  i<;o,s.  XVIII.,  p.  7. 


20 

n.     PRELIMINARY  EXPERIMENT. 

In  all  investigations  where  a  nitrogen  balance  is  attempted  it  is 
of  the  first  importance  to  bring  the  organism  to  be  used  as  sub- 
ject to  the  point  of  nitrogen  equilibrium  before  any  accurate  study 
of  the  specific  problem  shall  be  attempted.  To  this  end,  the  dog 
mentioned  on  page  19  was  selected  as  a  subject  and  preliminary 
feeding  of  the  animal  begun  on  Oct.  29,  1902.  During  the  first 
few  days  no  analyses  of  excreta  were  made  but  the  animal  was 
given  the  regulation  diet.  In  the  course  of  this  period  his  body 
weight  fell  from  17.23k.  on  Oct.  29  to  17.02k.  on  Nov.  i.  Be- 
ginning Nov.  2  full  analytical  data  were  collected. 

This  preliminary  experiment  embraced  a  period  of  twelve 
days.  In  that  time,  upon  a  regular  daily  ingestion  of  500  cc.  of 
water  the  dog  excreted  an  average  daily  volume  of  489  cc.  of 
urine.  Making  allowance  for  the  evaporation  which  must  of 
necessity  have  been  several  cubic  centimetres  daily  we  see  that 
practically  all  of  the  ingested  water  was  recovered.  It  will  be 
noted  that  the  body  weight  of  the  animal  was  wonderfully  regular 
during  the  early  days  of  the  experiment  and  even  at  the  twelfth 
day  showed  but  a  slight  decrease. 

For  this  preliminary  period  the  average  nitrogen  output  was 
9.673  grams  daily  (Table  III).  The  reaction  of  the  urine  was 
acid  throughout  the  period.  For  the  purpose  of  direct  compari- 
son with  other  data  the  last  five  days  of  the  experimental  period 
are  alone  taken  into  account.  Naturally  the  first  days  of  the  ex- 
periment when  the  dog  was  among  totally  new  surroundings  and 
being  fed  an  unaccustomed  diet,  the  collected  data  would  have 
comparatively  little  value.  However,  as  he  became  accustomed 
to  both  diet  and  conditions  the  organism  would  gradually  assume 
a  definite  nutritional  plane  and  nitrogen  equilibrium  would  result. 
Our  data  make  it  evident  that  from  the  eighth  to  the  twelfth  day, 
inclusive,  the  animal  was  rapidly  approaching  this  condition.  Dur- 
ing these  five  days  the  average  daily  volume  of  urine  was  517  cc. 
It  will  be  noted  that  the  average  urine  volume  for  the  first  seven 
days  was  but  468  cc,  indicating  an  evident  retention  of  water  by 
the  organism.  Therefore  the  increase  in  volume  during  the  fol- 
lowing five  days  was  to  have  been  expected.  During  this  same 
five  day  period,  upon  a  daily  nitrogen  ingestion  of  10.271  grams. 


21 

I0.023  grams  was  eliminated,  thus  showing  a  gain  of  but  0.248 
gram  daily  (Table  IV).  This  then  afforded  a  very  satisfactory 
starting  point  for  our  study  of  the  influence  of  hemorrhage. 

III.     FIRST  HEMORRHAGE 

In  determining  the  influence  of  the  withdrawal  of  blood  upon 
metabolism  it  is  in  many  ways  desirable  to  take  the  fluid  from  a 
small  vessel  supplying  a  comparatively  unimportant  area.  At 
first  thought  the  facial  vessels  seem  to  ofifer  many  advantages  in 
this  direction.  The  ligation  of  these  vessels,  taking  their  course 
as  they  do  along  boney  structures,  would  naturally  occasion  less 
disturbance  of  the  metabolic  activities  than  the  ligation  of  vessels 
of  the  same  calibre  in  many  other  parts  of  the  body.  It  was  as- 
certained, however,  by  means  of  a  preliminary  examination  of 
these  vessels  in  another  animal,  that  it  would  not  be  expedient  to 
attempt  to  draw  any  large  amount  of  blood  from  this  locality. 
Therefore,  after  due  consideration,  it  was  determined  to  make 
use  of  the  femoral  artery  somewhere  along  the  lower  portion  of 
its  course  (Saphenous  branch). 

I.  Operative  Procedure. — The  subject  of  this  experiment  was  a 
17k.  dog  which  had  been  on  a  fixed  diet  (See  page  55)  since  Oct. 
29  and  was  now,  after  sixteen  days  feeding,  approximately  in 
nitrogen  equilibrium  (See  Table  IV,  p.  58). 

In  order  to  make  impossible  the  loss  of  urine  should  any  be 
voided  during  anaesthesia,  the  beast  was  placed  in  a  large  zinc- 
lined  tray.  Urine  passed  under  these  conditions  could  easily  be 
recovered  by  means  of  a  pipette  and  filter  paper.  At  8.10  A.  M. 
the  administration  of  the  anaesthetic  was  begun.  From  the  very 
fir.st  the  beast  struggled  in  a  very  violent  manner,  necessitating 
the  combined  efforts  of  three  men  in  order  to  administer  the 
anaesthetic  at  all  satisfactorily.  At  the  end  of  eight  minutes,  as 
the  animal  still  possessed  much  of  his  original  vigor,  a  small 
amount  of  chloroform  (5  cc. )  was  added  to  the  ether  previously 
administered.  This  chloroform-ether  mixture  had  the  desired  ef- 
fect and  the  dog  was  soon  in  a  semi-unconscious  condition. 
Ether  was  now  continued  and  at  8.45  it  was  plainly  evident  that 
it  had  been  administered  to  the  surgical  degree.     The  ojieration 


22 

was  begun  at  this  time,  and  precautions  were  taken  throughout 
to  keep  the  conditions  strictly  asceptic. 

Before  laying  bare  and  clamping  the  artery  the  following 
arrangements  were  made :  A  cannula  previously  coated  on  the 
interior  with  an  extremely  thin  layer  of  mutton  tallow,  tO' 
prevent  the  blood  from  clotting,  was  adjustetd  to  a  rubber 
tube  of  appropriate  length  and  calibre.  In  collecting  the  blood  use 
was  made  of  a  25%  solution  of  sodium  chloride,  as  it  was  desir- 
able to  prevent  clotting  as  far  as  possible.  A  large  beaker  con- 
taining 273  grams  of  the  sodium  chloride  solution  (244  cc.)  was 
accurately  weighed,  after  which  both  balance  and  beaker  were 
carefully  supported  in  a  position  very  near  and  somewhat  below 
the  operating  table.  The  balance  was  placed  in  this  position  in 
order  to  make  the  length  of  tube  leading  from  the  cannula  to  the 
beaker  as  short  as  possible  and  in  this  way  further  guard  against 
clotting.  It  had  previously  been  determined  to  withdraw  an 
amount  of  blood  approximating  3%  of  the  animal's  body  weight, 
or  about  500  grams.  To  accomplish  this  accurately  a  500  gram 
weight  was  added  to  the  weights  of  the  beaker  and  salt  solution 
and  it  was  then  arranged  to  allow  the  hemorrhage  to  continue 
until  it  was  evident  from  the  movement  of  the  balance  pan  that 
approximately  the  desired  amount  had  been  collected.  As  soon 
as  possible  after  the  completion  of  the  above  arrangements  the 
artery  was  laid  bare  and  the  blood  drawn. 

In  forty-six  minutes  from  the  time  the  anaesthetic  was  discon- 
tinued the  dog  was  entirely  himself,  stood  on  his  feet,  and  al- 
though he  was  somewhat  weak  from  loss  of  blood  and  the  effect 
of  the  anaesthesia,  he  appeared  in  fine  condition.  At  no  time  after 
the  operation  was  there  any  inc.lmation  to  vomit,  and  the  appetite 
of  the  animal  continued  excellent  throughout  the  period. 

Time  Schedule. — 8.10  A.  M.,  anaesthesia  begun  (ether).  8.18, 
trace  of  chloroform  given.  8.35,  beast  placed  on  board.  8.45, 
operation  begun.  9.26,  incisions  made,  artery  laid  bare,  and 
ligated,  cannula  inserted  and  slow  hemorrhage  commenced. 
10.00,  commencement  of  more  rapid  hemorrhage.  10.12,  return 
to  slow  hemorrhage.  10.14,  hemorrhage  ended.  10.20,  wounds 
sewed  up  and  administration  of  ether  discontinued.  10.21,  beast 
removed  from  board  to  cage.     10.25,  evidence  of  returning  con- 


23 

sciousness.  10.55,  sat  up  in  his  cage.  11.06,  stood  on  his  feet 
and  moved  about.  The  blood  drawn  at  this  hemorrhage  weighed 
492.9  grams  (Table  II). 

2.  Observations  on  the  Influence  of  the  First  Hemorrhage. — 
The  first  observation  following  the  hemorrhage  was  the  failure 
of  the  dog  to  urinate  as  frequenth'  or  as  large  a  volume  as  former- 
ly. The  experiment  started  at  8.10  A.  M.  and  the  first  urine  was 
not  passed  until  from  16  to  24  hours  afterward  (Table  V).  The 
urine  volume  for  the  day  of  the  operation  was  therefore  the  low- 
est of  any  day  of  the  investigation  up  to  that  time.  Now  in  our 
control  experiments  we  have  seen  that  under  the  influence  of  the 
anaesthesia,  in  each  case,  the  first  urine  was  passed  in  from  7  to 
8  hours  after  the  operation.  It  is  also  seen  from  our  data  that 
on  the  first  day  of  the  control  experiments  the  maximum  volume 
of  urine  was  eliminated.  Therefore  this  very  low  volume  after 
hemorrhage  is  all  the  more  significant  when  we  consider  that  the 
same  force  (anaesthesia)  was  acting  at  the  time  of  the  hemor- 
rhage as  was  at  work  when  we  secured  the  very  large  urine 
elimination  on  the  initial  days  of  our  control  experiments.  The 
onlv  new  factor  is  the  iiemorrhage.  Hence  the  influence  of  the 
withdrawal  of  blood  is  much  greater  than  appears,  for  in  order 
to  so  delay  the  flow  of  urine  that  the  first  elimination  should  not 
occur  until  from  16  to  24  hours  after,  the  hemorrhage  has  been 
forced  to  overcome  the  diuretic  influence  of  the  anaesthesia  which 
would  have  caused  a  large  output  of  urine  in  from  7  to  8  hours 
after. 

The  nitrogen  content  (6.33  grams)  of  the  urine  on  the  first  day, 
was  also  the  minimum  outi)ut  up  to  date.  The  body  weight  of  the 
dog  fell  0.51k.  on  the  first  day.  On  the  second  day  some  radical 
changes  occurred.  On  this  day  the  dog  passed  an  unusually  large 
volume  of  urine,  having  the  maximum  specific  gravity  for  the  ex- 
periment, and  containing  the  second  largest  content  of  nitrogen. 
The  body  weight  of  the  dog  continued  to  fall  somewhat. 

Notwithstanding  the  rather  high  volumes  of  urine  passed  on  the 
second  and  third  days  after  the  hemorrhage,  the  average  at  any 
time  was  considerably  below  that  of  the  preliminary  experiment. 
For  instance,  the  average  of  the  first  nine  days  was  but  467  cc, 
whereas  the  average  preceding  the  hemorrhage  was  489  cc.    This 


24 

showed  an  average  retention  by  the  organism  of  22  grams  of 
water  daily,  or  198  grams  in  the  nine  days.  During  the  last  seven 
days  of  the  experiment  the  retained  water  seemed  to  have  been 
eliminated  and  the  normal  ratio  again  reached.  In  the  course 
of  these  seven  days  the  average  urine  volume  was  518  cc,  a  daily 
increase  of  29  cc.  above  that  of  the  preliminary  experiment,  or  a 
total  increase  of  203  grams.  From  the  weights  as  determined  we 
find  that  during  this  period  of  seven  days  the  dog's  weight  fell 
from  16.34k.  to  16.09k.,  a  total  loss  of  250  grams. 

3.  Discussion  of  the  Income  and  Outgo  of  Nitrogen,  Sulphur  and 
Phosphorus  After  the  First  Hemorrhage. — The  study  of  the  first 
hemorrhage  was  begun  with  the  subject  showing  a  gain  of  0.248 
gram  of  nitrogen  daily.  The  effect  of  the  withdrawal  of  blood 
was  to  cause  an  increase  in  the  nitrogen  content  of  the  urine,  this 
increase  appearing  the  first  day  after  the  hemorrhage,  and  reach- 
ing its  maximum  at  the  fifth  day.  After  this  date  the  average 
elimination  decreased  and  approximate  nitrogen  equilibrium  was 
reached  on  the  sixteenth  day  of  the  experiment. 

By  referring  to  the  data  for  the  nitrogen  elimination  for  the  five 
days  following  this  hemorrhage  (Table  IV),  it  will  be  seen  that 
the  average  daily  increase  in  nitrogen  for  this  period  was  1.009 
grams.  The  dog  was  gaining  0.248  gram  of  nitrogen  daily,  how- 
ever, at  the  time  of  the  hemorrhage,  hence  the  effect  was  to  cause 
an  increased  output  of  1.257  grams  of  nitrogen  daily.  This  rep- 
resents the  final  effect  produced  by  the  anaesthesia,  operation  and 
hemorrhage  acting  in  unison.  Now  we  shall  see  from  our  con- 
trol experiments  (Table  XIX),  that  the  effect  of  the  anaesthesia 
was  to  decrease  the  nitrogen  output  0.937  gram  daily.  Making 
this  correction  we  have  as  the  effect  of  operation  and  hemorrhage 
an  increase  of  2.194  grams  daily,  and  further  correcting  for  the 
effect  of  the  operation  we  have  as  the  net  effect  of  the  hemorrhage 
an  increase  in  the  daily  nitrogen  output  of  0.488  gram  (Table 
XIX). 

The  nitrogen  balance  for  the  whole  sixteen  days  of  the  experi- 
ment (Table  III)  showed  a  loss  of  0.179  gram  of  nitrogen  daily. 
The  sulphur  followed  this  with  a  daily  loss  of  0.023  gram  (Tables 
VI  and  XI)  while  the  phosphorus  showed  a  daily  gain  of  0.123 
grams. 


25 

rV.     CONTROL  EXPERIMENTS. 

In  studying-  the  effect  of  the  withdrawal  of  blood  there 
were  evidently  two  forms  of  control  experiments  necessary 
before  any  definite  conclusions  could  be  reached.  The  first 
of  these  was  the  influence  of  anaesthesia,  and  the  sec- 
ond the  influence  of  the  anaesthesia  plus  the  influence 
of  the  operation  which  accompanied  all  our  withdrawals 
of  blood.  If  it  were  practicable  to  draw  blood  without  the 
use  of  any  anaesthetic,  obviously  the  first  form  of  control  ex- 
periment could  be  eliminated.  But  even  were  such  a  method  pur- 
sued other  factors,  such  as  the  nefvous  influence,  would  be  intro- 
duced and  we  could  not  be  certain  that  the  effect  secured  was  de- 
pendent alone  upon  loss  of  blood.  Obviously,  for  this  reason  ex- 
periments upon  the  influence  of  hemorrhage,  made  withotit  the 
use  of  anaesthetics,  would  be  extreiiiely  difficult  to  control.  Local 
anaesthetics  might  be  employed,  but  even  here  we  could  not  be 
sure  what  influence  tlie  absorption  of  these  bodies  w-ould  have 
upon  the  organism.  The  experimental  plan  adopted  by  us  was 
open  to  complete  control  which  we  feel  would  not  have  been  possi- 
ble had  we  followed  either  of  the  other  plans  mentioned. 

Two  control  experiments  were  made,  the  first,  ten  days  in 
length,  was  devoted  to  the  study  of  the  influence  of  anaesthesia, 
and  in  the  second,  covering  a  period  of  thirteen  days,  an  attempt 
was  made  to  determine  the  effect  of  anaesthesia  accompanied  by 
the  operative  manipulation. 

I.  Influence  of  Anaesthesia. — The  general  procedure  in  this 
our  first  control  experiment  was  practically  identical  with  that  of 
the  regular  blood-letting  experiments.  The  animal  was  placed  on 
the  operating  board,  loosely  tied  to  allow  freedom  of  motion  in 
his  struggles,  and  the  administration  of  ether  begun.  After  an  in- 
terval of  a  few  moments  a  small  amount  of  chloroform  was  given 
as  in  the  other  experiments.  Care  was  taken  to  consume  the  usual 
length  of  time  in  reaching  complete  anaesthesia.  The  beast  was 
kept  under  the  influence  of  the  anaesthetic  for  a  period  corre- 
sponding to  the  normal  time  of  operation  and  was  then  returned 
to  his  cage.  A  detailed  time  schedule  follows: — 9.10,  anaesthesia 
begun.     9.18,  chloroform  administered.     9.40,  anaesthesia  com- 


26 

plete.  1 1. 20,  administration  of  ether  discontinued  and  dog  re- 
turned to  cage.  11.30,  returning  consciousness.  11.50,  sits  up 
and  staggers  about.  12.00,  appears  normal.  The  whole  period  of 
anaesthesia  was  without  incident  except  the  loss  of  about  5  cc. 
of  urine  at  9.15  by  reason  of  vigorous  abdominal  movement.  This 
urine  was  recovered  by  means  of  filter  paper  and  added  to  the 
washings  for  the  period.  The  struggling  incident  to  the  former 
anaesthesia  was  duplicated  in  detail. 

(a).     Observations  on  the  Influence  of  Anaesthesia. 

Before  attempting  to  study  the  influence  of  anaesthesia  an  ef- 
fort was  made  to  get  the  dog  as  nearly  as  possible  in  a  condition  of 
nitrogen  equilibrium.  By  referring  to  the  nitrogen  balance  for 
the  five  days  immediately  preceding  the  anaesthesia  experiment 
(Table  IV,  p.  58)  it  will  be  seen  that  the  animal  was  approximate- 
ly in  the  desired  condition,  there  being  a  loss  of  but  0.159  gram 
of  nitrogen  per  day. 

Upon  an  examination  of  the  data  for  this  experiment  (Table 
XII,  p.  66)  several  evident  variations  from  the  order  ob- 
taining in  the  preliminary  and  first  blood-letting  experiments 
are  noted.  Whereas  the  average  urine  volume  of  the  pre- 
ceeding  periods  had  been  489  cc.  in  each  case,  here,  upon 
the  same  ingestion  of  water  we  secured  an  average  urine 
volume  of  522  cc.  for  the  period.  The  urine  volume  for 
the  day  on  which  the  anaesthetic  was  administered  was 
particularly  worthy  of  notice.  Upon  this  day,  which  in 
the  experiment  preceeding  had  shown  a  urine  volume  of  377 
cc.  we  secured  a  volume  of  654  cc,  or  nearly  double  that  pre- 
viously secured.  When  we  compare  this  volume  with  the  average 
volume  for  the  whole  period  preceeding  we  still  have  an  increase 
of  165  cc.  for  the  day.  This  increase  in  volume  is  evidently 
traceable  to  the  diuretic  action  of  the  ether. 

Another  striking  feature  of  the  urine  excretion  was  its  high 
specific  gravity  accompanied  by  a  relatively  low  nitrogen  content. 
This  was  especially  true  of  the  first  few  days  following  anaes- 
thesia. As  measured  by  other  daily  excretions,  both  before  and 
after  the  anaesthesia  experiment,  a  urine  volume  of  654  cc.  hav- 
ing a  specific  gravity  of  1.018  would  have  given  us  a  nitrogen  out- 
put of  10.94  grams  (See  data  for  Nov.  10,  Nov.  24,  Jan.  12,  and 


27 

Tan.  13).     Tims  the  effect  of  the  anaesthesia  was  to  diminish  the 
total  nitrogen  excretion  for  the  day  approximately  20%. 

After  the  excretion  of  phosphates  and  sulphates  failed  to  show 
the  cause  of  the  unusual  relation  between  the  specific  gravity  and 
nitrogen  content  we  turned  to  the  chlorides  for  a  solution  of  the 
problem,  and  beginning  Nov.  25  duplicate  determinations  of  the 
chlorine  content  of  the  urine  were  made  for  every  day  up  to  and 
including  Dec.  3.     The  following  results  were  obtained : — 

Xov.  25 1.82  grams  of  chlorine. 

Nov.  26 1. 71  grams  of  chlorine. 

Nov.  2"] 1.65  grams  of  chlorine. 

Nov.  28 1.87  grams  of  chlorine. 

Nov.  29 1.69  grams  of  chlorine. 

Nov.  50 ^.J7  grams  of  chlorine. 

Dec.     I 2.85  grams  of  chlorine. 

Dec.     2 2.73  grams  of  chlorine. 

Dec.     3 2.39  grams  of  chlorine. 

From  this  data  it  will  be  seen  that  the  chlorine  output  was  fair- 
Iv  constant  up  to  the  day  the  anaesthetic  was  given,  showing  an 
average  daily  elimination  of  1.75  grams,  and  that  on  this  day  an 
increase  of  150%  in  the  chlorine  content  of  the  urine  occurred. 
Thus  the  indications  are  that  the  rise  in  the  daily  output  of  chlor- 
ides through  the  influence  of  anaesthesia,  caused  a  higher  specific 
gravity  on  the  days  immediately  following  the  administration  of 
the  anaesthetic  than  the  determined  content  of  nitrogen  would 
warrant. 

As  a  further  verification  we  took  a  volume  of  water  (649  cc.) 
equal  to  the  volume  of  urine  passed  by  the  dog  on  Nov.  30,  and 
dissolved  in  this  7.2  grams  of  sodium  chloride,  which,  according 
to  our  data,  was  the  amount  eliminated  on  that  day.  This  gave 
us  a  specific  gravity  of  1.009.  T'le  actual  specific  gravity  of  the 
urine  of  that  day  being  1.018,  we  see  that  a  large  part  of  the  den- 
sity of  the  fluid  was  due  to  the  presence  of  sodium  chloride.  We 
next  added  to  this  sodium  chloride  solution  an  amount  of  urea 
C18.4  grams)  equivalent  to  the  nitrogen  content  of  the  urine  for 
that  day,  and  secured  a  specific  gravity  of  1.015.  This  left  but 
three  points  in  density  to  be  met  by  the  other  solid  ingredients  of 


28 

the  urine.  Thus  we  secured  from  7.2  grams  of  sodium  chloride  a 
rise  in  specific  gravity  50%  greater  than  from  18.4  grams  of  urea. 
This  rough  calculation  easily  explained  the  condition  of  high  spe- 
cific gravity  and  low  nitrogen  content. 

As  has  been  said  elsewhere  (p.  23)  the  drawing  of  blood, 
among  other  well  marked  effects,  caused  a  very  noticeable  de- 
crease in  the  amount  of  urine  eliminated  for  an  extended  period 
thereafter.  Consider,  for  example,  the  hemorrhages  immediately 
preceeding  and  following  the  anaesthesia  experiment.  After  the 
first  hemorrhage  which  began  at  8.10  A.  M.  no  urine  was  passed 
during  the  next  sixteen  hours,  or  up  to  midnight  (Table  V).  On 
the  following  morning  at  8.30  the  urine  volume  was  300  cc.  As 
the  beast  was  not  observed  between  midnight  and  8.30  A.  M.  we 
are  unable  to  conclude  at  what  hour  this  urine  was  eliminated. 
After  the  second  hemorrhage  a  similar  course  was  followed,  i.  e., 
no  urine  up  to  midnight,  followed  by  a  volume  of  590  cc.  at  9 
o'clock  the  next  morning.  In  the  anaesthesia  experiment,  how- 
ever, where  no  blood  was  drawn,  but  the  individual  influence  of 
the  anaesthetic  was  studied,  we  find  very  different  conditions  to 
prevail.  Here,  following  the  administration  of  the  ether  at  9.10 
A.  M.,  we  observed  a  fairly  large  elimination  of  urine  (283  cc.) 
in  seven  hours  (4.15  P.  M.)  or  in  less  than  one-half  the  time  that 
elapsed  before  the  first  urine  appeared  after  the  hemorrhages 
mentioned.  The  diuretic  influence  of  the  ether  was  thus  very 
clearly  indicated. 

As  was  observed  in  the  other  periods,  the  body  weight  of  the 
animal  fell  somewhat  upon  the  day  of  anaesthesia,  in  this  particu- 
lar casea  loss  of  0.28k.  being  noted. 

(b).    Discussion  of  the  Income  and  Outgo  of  Nitrogen,  Sulphur  and 
Phosphorus  Before  and  After  Anaesthesia. 

It  is  very  evident  that  an  effect  produced  by  an  agency  such  as 
anaesthesia  or  hemorrhage  will  naturally  be  more  pronounced 
during  the  days  immediately  following  the  operation.  It  is  also 
easily  seen  that  in  all  such  instances  there  will  come  a  point,  soon- 
er or  later,  at  which  time  the  animal  will  again  begin  to  approach 
the  normal  and  tend  toward  nitrogen  equilibrium.  Such  being 
the  situation  it  is  of  course  essential  to  know  as  nearly  as  possible 


29 

when  tiitrogen  equilibrium  is  being  reached.  In  order  to 
facihtate  comparison  and  the  drawing  of  conckisions,  ni- 
trogen balances  for  a  short  period  immediately  preceed- 
ing  the  commencement  of  each  experiment  have  been 
made  in  order  to  show  the  exact  state  of  the  organ- 
ism at  the  moment  the  experiment  began.  In  addition,  nitro- 
gen balances  for  a  few  days  following  the  commencement  of  each 
experiment  are  given  in  order  to  show  the  immediate  effect,  if 
anv.  which  was  secured  in  each  specific  instance.  No  attempt  be- 
ing made  after  Jan.  19  to  get  the  dog  again  into  nitrogen  equili- 
brium, such  balances  will  not  be  found  for  experimental  days  after 
that  date.  As  sulphur  and  phosphorus  were  not  determined  in 
daily  samples  no  such  balances  will  be  found  for  these  elements. 

By  referring  to  the  nitrogen  balance  for  the  short  period  imme- 
diately preceeding  the  administration  of  the  anaesthetic  (Table 
IV }  it  will  be  seen  that  the  dog  was  approximately  in  nitrogen 
equilibrium,  and  by  consulting  this  table  further  we  note  that  the 
effect  of  anaesthesia  was  to  cause  a  lowering  of  the  nitrogen  out- 
put and  a  consequent  daily  gain  of  0.937  gram  of  nitrogen  to  the 
body.  Data  concerning  the  relation  between  the  nitrogen  content 
and  the  specific  gravity  of  the  urine  during  this  period  will  be 
found  on  page  26. 

The  course  of  the  sulphur  excretion  in  a  general  way  followed 
that  of  the  nitrogen.  The  period  preceeding  the  anaesthesia  ex- 
periment showed  a  daily  loss  of  0.023  gram  of  sulphur  (Table 
VI),  whereas  the  balance  for  the  period  showing  the  influence 
of  the  anaesthetic  exhibited  a  gain  of  0.015  gram  of  sulphur 
daily.  Thus  we  had  as  the  final  effect  of  the  anaesthesia  upon  the 
course  of  the  sulphur  excretion  a  daily  gain  of  0.128  gram. 

The  course  of  the  phosphorus  excretion  seemed  to  be  directly 
opposite  that  of  nitrogen  and  sulphur.  Showing  before  the  ex- 
periment a  gain  of  0.123  gram  of  phosphorus  daily  (Table  VII), 
we  secured  after  the  anaesthesia  a  gain  of  but  0.015  gram  daily. 
In  this  instance  the  ])hosphorus  showed  a  loss  of  0.108  gram 
daily,  whereas  the  nitrogen  and  sulphur  each  showed  a  gain.  Thus 
the  anae.sthsia  caused  a  fall  in  the  daily  excretion  of  nitrogen  and 
sulphur  anrl  a  coincident  rise  in  the  phosphorus  excretion. 

2.  Influence  of  Anaesthesia-Operation. — The  nitrogen  content 


30 

of  the  urines  of  Dec.  6-9  indicating  that  the  dog  was  approximate- 
ly in  nitrogen  equiHbrium  (-|- 0.158  gram  of  nitrogen  per  day) 
it  was  determined  to  begin  our  second  control  experiment  on  Dec. 
10.  Apart  from  the  drawing  of  blood,  the  procedure  in  this  ex- 
periment followed  the  same  course  as  that  of  the  regulation  blood- 
letting experiments.  The  beast  was  anaesthetized  and  the  custom- 
ary operative  steps  were  carefully  followed.  The  regulation  incis- 
ion was  made,  artery  laid  bare,  (in  this  case  the  right  femoral-) 
clamped  and  the  cannula  inserted.  The  wound  was  then  sewed  up 
and  the  animal  returned  to  his  cage.  In  fact  everything  was  done 
to  make  this  operation  coincide  in  every  detail  with  the  regulation 
blood-letting  operations.  Schedule  follows: — 8.12  A.  M.  anaes- 
thesia begun. ^  8.20,  small  amount  of  chloroform  given.  8.45, 
operation  begun.  10.23,  anaesthetic  discontinued.  10.24,  dog 
placed  in  cage.  10.28,  signs  of  returning  consciousness.  10.  40, 
sits  up. 

(a).     Observations  on  the  Influence  of  Anaesthesia-Operation. 

In  the  main  the  characteristics  of  this  period  were  similar  to 
those  observed  during  the  anaesthesia  experiment.  Here  with  the 
same  uniform  water  ingestion  of  500  cc.  we  secured  an  average 
daily  urine  volume  of  531  cc,  thus  comparing  favorably  with  the 
average  (522  cc.)  of  the  anaesthesia  period,  but  being  in  marked 
contrast  to  the  average  elimination  (489  cc.)  secured  during  our 
preliminary  experiment. 

The  same  condition  of  low  nitrogen  elimination,  when  ap- 
proached from  the  point  of  view  of  specific  gravity,  maintained 
here  as  was  seen  to  prevail  during  the  early  part  of  the  anaesthesia 
experiment.  Here  as  in  the  former  instance  we  considered  that 
a  largely  increased  output  of  chlorine  caused  the  peculiar  rela- 
tion. 

In  this  experiment,  as  well  as  in  the  one  devoted  exclusively  to 
the  study  of  the  influence  of  ether,  we  noted  a  copious  flow  of 
urine  appearing  much  earlier  than  after  the  blood-letting  opera- 
tions. As  in  the  anaesthesia  experiment,  this  copious  flow  of 
urine  was  doubtless  due  to  diuresis  produced  by  the  anaesthetic. 

On  Dec.  18  at  about  i  P.  M.  the  dog  was  observed  lapping  its 

IThe  animal  passed  a  few  grams  of  feces  during  preliminary  anaesthesia.     This  waS 
added  to  the  feces  for  the  day. 


31 

wound,  and  upon  examination  it  was  found  that  a  slight  hemor- 
rhage had  been  produced  by  the  animal,  probably  by  means  of  his 
foot  in  the  act  of  scratching.  The  flow  of  blood  was  soon  stopped, 
however,  and  the  wound  bandaged  with  adhesive  plaster  to  pre- 
vent the  animal  producing  another  hemorrhage.  Probably  not 
over  25  cc.  of  blood  was  lost  and  practically  all  of  this  was  eaten 
by  the  dog.  By  an  examination  of  the  data  it  will  be  seen  that  the 
nitrogen  elimination  of  Dec.  18  was  the  highest  of  the  experiment, 
while  that  of  Dec.  19  was  also  relatively  high.  This  increase  may 
have  been  due  partly  to  the  ingested  blood,  or  it  may  have  been  the 
normal  excretion  since  these  two  days  were  followed  by  two  days 
of  low  elimination  which  upon  the  21st  practically  restored  the 
ratio  as  it  existed  previous  to  the  accidental  hemorrhage. 

(b).    Discussion  of  the  Income  and  Outgo  of  Nitrogen,  Sulphur  and 
Phosphorus  Before  and  After  Anaesthesia-Operation. 

The  nitrogen  balance  for  the  four  days  immediately  preceeding 
the  commencement  of  the  anaesthesia-operation  experiment  (Table 
IV)  showed  us  that  the  dog  was  practically  in  a  condition  of  ni- 
trogen equilibrium  (-[-0.158  gram  daily).  The  next  balance, 
i.  €..  that  for  the  five  days  immediately  following  the  anaesthesia- 
operation  of  Dec.  10,  records  an  average  daily  loss  of  0.61 1  gram 
of  nitrogen.  Hence  the  efifect  of  the  anaesthesia-operation  was 
to  cause  an  increase  of  0.769  gram  in  the  daily  nitrogen  output 
(Table  XIX;.  Now  we  have  already  seen  that  the  effect  of  the 
anaesthesia  alone  was  to  decrease  the  nitrogen  elimination  0.937 
gram  daily.  Therefore  the  operation  alone  caused  an  increase 
in  the  nitrogen  output  large  enough  to  overcome  entirely  the  de- 
crease due  to  the  anaesthesia  and  yet  show  a  daily  increase  of 
0.769  gram.  Hence  the  total  increase  due  to  the  operation  was 
1.706  grams  daily  (Table  XIX). 

As  was  noted  in  the  anaesthesia  experiment  the  course  of  the 
sulphur  excretion  was  very  similar  to  that  of  the  nitrogen.  In 
the  case  of  the  phosphorus  excretion  also  a  uniformity  with  the 
conditions  obtaining  in  the  anaesthesia  experiment  was  noted. 
Tims  while  the  nitrogen  and  sulphur  showed  an  increased  daily 
elimination  as  a  result  of  anaesthesia-operation  the  phosphorus 
excretion  showed  a  decrease. 


32 

V.     SECOND  HEMORRHAGE. 

This  hemorrhage  occurred  on  Dec.  23  at  the  close  of  the  anaes- 
thesia-operation experiment.  An  attempt  was  again  made  ta 
withdraw  approximately  3%  of  the  animal's  body  weight.  Apart 
from  minor  details  the  operative  procedure  was  the  same  as  that 
observed  at  the  first  hemorrhage  (See  p.  21).  Two  incisions 
were  made  as  at  the  first  operation.  It  was  attempted  to  draw 
the  blood  from  a  branch  of  the  left  femoral  just  above  the  knee, 
but  the  vessel  being  too  small  at  this  point  a  second  incision  was 
made  higher  up  and  the  main  trunk  of  the  femoral  laid  bare.  A 
detailed  schedule  follows: — 8.14  A.  M.,  anaesthesia  begun.  8.20, 
trace  of  chloroform.  8.40,  first  incision  made.  9.12,  second  in- 
cision made.  9.30,  cannula  inserted.  9.31,  hemorrhage  begun. 
9.40,  hemorrhage  ended.  TO.05,  anaesthetic  discontinued.  10.10, 
wounds  sewed  up.  10.12,  animal  in  cage.  10.24,  returning  con- 
sciousness.    11.40,  dog  appears  normal. 

The  total  amount  of  blood  lost  to  the  organism  at  this  hemor- 
rhage was  506  grams. 

I.  Observations  on  the  Influence  of  the  Second  Hemorrhage. — 
According  to  Hammarsten^  a  hemorrhage  amounting  to  one- 
fourth  of  the  total  blood  produces  no  continued  sink- 
ing of  the  blood  pressure  in  the  arteries ;  a  loss  of 
one-third  reduces  the  blood  pressure  considerably  and  a 
loss  of  one-half,  in  adults,  is  dangerous  to  life.  Now 
in  our  second  hemorrhage  on  Dec.  23  we  drew  506 
grams  of  blood  from  the  animal  weighing  at  that  time  15.72k. 
Taking  the  amoimt  of  blood  present  as  being  ^/is.g  of  his  body 
weight  (Hammarsten's  figure)  or  1164  grams,  it  is  seen  that,  ac- 
cording to  Hammarsten,  a  hemorrhage  of  582  cc.  would  be  dan- 
gerous to  life.  The  hemorrhage  we  produced  amounted  to  506 
cc,  and,  as  our  data  shows,  the  dog  appeared  as  usual  two  hours 
afterward.  It  seems  hardly  probable  that  the  animal's  condition 
would  have  been  materially  different  had  we  drawn  76  cc.  more 
blood,  thus  securing  Y^  of  his  entire  supply. 

In  the  main  the  influence  of  the  second  hemorrhage  seemed  to 
be  closely  parallel  to  that  of  the  first  hemorrhage.  At  this  second 
hemorrhage  we  noted  the  low  volume  of  urine  (428  cc.)  on  the 

1  Phj'siological  Chemistry,  Fourth  Edition  (English),  1904,  p.  209. 


33 

day  blood  was  drawn,  followed  by  a  period  of  16-25  liours  (Table 
\')  during  which  no  urine  was  passed;  two  points  in  close  agree- 
ment with  those  of  the  first  hemorrhage.  The  urine  volume  ran 
much  higher  after  the  second  hemorrhage  than  after  the  first.  In 
the  four  days  following  the  second  hemorrhage  the  average  urine 
volume  was  554  cc.  (Table  XIl),  while  the  average  daily  elimina- 
tion for  seven  days  was  525  cc.  and  for  the  whole  21  days  of  the 
experiment  was  530  cc.  When  contrasted  with  the  average  of 
451  cc.  for  the  four  days  following  the  first  hemorrhage  and 
489  cc,  the  average  for  the  whole  experiment,  the  variations  are 
quite  impressive.  These  vohunes  seem  to  indicate  that  the  second 
hemorrhage  was  not  so  successful  in  masking  the  diuretic  effect 
of  the  anaesthesia  as  was  the  first  hemorrhage. 

The  body  weight  of  the  animal  fell  from  15.72k.  to  15.18k.  on 
the  first  day  of  the  experiment,  showing  a  loss  of  0.54k.  thus  very 
closely  agreeing  with  the  loss  of  0.51k.  sustained  through  the 
agency  of  the  first  hemorrhage.  The  body  weight  continued  to 
fall  until,  on  the  fourth  day,  a  weight  of  14.76k  was  registered. 
This  weight  was  maintained  practically  unaltered  for  six  days 
when  another  gradual  decrease  began  and  continued  until  the  end 
of  the  experiment.  The  total  loss  of  weight  in  the  twenty-one 
days,  exclusive  of  the  loss  due  to  hemorrhage,  was  0.76k.  The 
reaction  of  the  urine  was  as  a  rule  amphoteric  during  the  early 
days  of  the  experiment,  the  alkalinity  gradually  decreasing  until, 
during  the  closing  days  of  the  expepiment  the  amphoteric  charac- 
ter had  disappeared  and  an  acid  reaction  obtained. 

In  agreement  with  the  observations  following  the  first  hemor- 
rhage the  second  day  showed  us  a  large  volume  of  urine  having 
the  highest  specific  gravity  of  any  urine  of  the  experiment,  and 
likewise  containing  the  highest  nitrogen  content. 

2.  Discussion  of  the  Income  and  Outgo  of  Nitrogen,  Sulphur  and 
Phosphorus  After  the  Second  Hemorrhage. — This  experiment  start- 
ed with  the  dog  ap];roximately  at  nitrogen  equilibrium,  ;'.  c,  show- 
ing daily  loss  of  but  0.141  gram  of  nitrogen.  The  loss  of  blood 
caused  this  condition  to  rapidly  change  into  one  of  largely  in- 
creased nitrogen  elimination  and  at  the  end  of  the  seventh  day  a 
daily  increase  of  1.799  R^ams  of  nitrogen  was  noted.  Making  the 
projR-r  correction  for  the  influence  of  the  anaesthetic  (See  p.  75) 


34 

we  have,  for  the  first  seven  days  a  dail}^  increase  of  2.736  grams  of 
nitrogen  as  the  combined  effect  of  the  operation  and  hemorrhage, 
and  correcting  further  for  the  operation,  we  obtain  a  daily  in- 
crease of  1.03  grams  as  the  individual  effect  of  the  hemorrhage 
(Table  XIX).  It  will  be  recognized  that  this  increase  was  some- 
what higher  than  the  increase  due  to  the  first  hemorrhage.  The 
larger  urine  volume  may  have  had  some  influence  in  this  direction. 
The  last  five  days  of  the  experiment  indicate  a  loss  of  0.150  gram 
of  nitrogen  daily,  comparing  very  favorably  witih  the  average  loss 
of  0.141  gram  at  the  beginning. 

In  agreement  with  the  large  increase  in  the  nitrogen  excretion 
following  hemorrhage,  the  excretion  of  sulphur  was  also  un- 
usually high.  This  element  showed  an  increase  of  0.180  gram 
per  day  for  the  entire  experiment  (Table  VI).  The  excretion  of 
phosphorus,  as  usual,  followed  a  course  typically  its  own  and  in- 
stead of  showing  an  increase  as  did  the  nitrogen  and  sulphur,  it 
registered  a  decreased  excretion  amounting  to  0.163  gram  per  day 
(Table  VII). 

VI.     THIRD  HEMORRHAGE. 

The  third  hemorrhage  occurred  on  the  morning  of  Jan.  13,  just 
21  days  after  the  second  hemorrhage.  The  dog  at  this  time  was 
seemingly  in  fine  condition  and  was  using  the  leg,  operated  upon 
at  the  second  hemorrhage,  in  a  normal  manner.  It  was  decided  to 
draw  blood  on  this  occasion  from  the  left  femoral  artery  slightly 
higher  up  than  the  site  of  the  second  hemorrhage. 

The  operation  began  at  8.20  A.  M.,  the  technique  being  analo- 
gous to  that  of  former  operations.  Upon  making  the  incision  we 
were  much  surprised  at  the  altered  position  of  the  femoral  artery. 
Down  beneath  and  around  the  wound  of  the  second  hemorrhage 
the  central  end  of  the  artery  was  surrounded  by  a  peculiar  cap- 
sule formation  which  pushed  upon  the  artery  causing  it  to  lie 
deeply  imbedded  among  the  tissues.  After  inserting  the  cannula 
the  hemorrhage  was  allowed  to  continue  ten  minutes  at  the  end  of 
which  period  it  was  evident  from  physical  signs  that  it  would  not 
be  policy  to  continue  the  hemorrhage  longer.  The  tongue  of  the 
beast  was  much  more  blanched  than  at  previous  hemorrhages  and 
the  pulse  in  the  right  femoral  was  almost  imperceptible.       The 


35 

animal  was  so  very  weak  that  the  sewing  up  of  the  wound  was 
hastened  and  the  administration  of  ether  discontinued  as  soon  as 
possible.  Thus  the  period  of  anaesthesia  was  somewhat  shorter 
than  at  the  previous  operations.  The  wound  of  the  second  hem- 
orrhage was  drained,  a  few^  deep  lying  stitches  were  removed,  and 
the  whole  wound  cleansed  with  carbolic  acid.  A  schedule  of  the 
operative  steps  follows : — 8.20  A.  M.,  anaesthesia  begun.  8.28, 
trace  of  chloroform.  8.41,  operation  begun.  9.14,  cannula  insert- 
ed. 9.15,  hemorrhage  begun.  9.20,  ether  discontinued.  9.24, 
hemorrhage  ended.  9.52.  wound  sewed  up.  9.53,  animal  in  cage. 
10.02.  returning  consciousness.     10.20,  dog  sits  up. 

There  was  somewhat  less  salivation  than  at  previous  operations. 
A  short  time  after  the  operation  the  dog  used  his  leg  much  more 
handily  than  after  former  hemorrhages.  The  total  blood  lost  to 
the  organism  aggregated  505.5  grams. 

I.  Observations  on  the  Influence  of  the  Third  Hemorrhage. — A 
larger  percentage  of  the  total  blood  of  the  animal  was  drawn  at 
this  hemorrhage  than  upon  any  other  occasion  during  the  course 
of  the  experiments.  As  has  been  said  the  blood  was  allowed  to 
flow  until  the  animal  was  in  an  extremely  weakened  state.  When 
the  weights  were  made  it  w-as  found  that  the  total  hemorrhage 
had  been  505.5  grams  or  3.51%  of  the  body  weight  of  the  animal. 
According  to  the  theory  of  Hammarsten  (loc.  cit.)  a  hemorrhage 
of  534  grams  would  have  been  fatal.  We  are  inclined  to  believe 
that  a  short  continuation  of  the  hemorrhage  would  have  been  at- 
tended by  fatal  results ;  but,  in  view  of  the  rapid  recovery  of  the 
dog  we  do  not  believe  that  an  increase  of  28.5  grams  in  the 
amount  of  blood  drawn  would  have  caused  the  death  of  the  ani- 
mal. It  was  very  evident,  how^ever,  that  the  dog  was  much  more 
effected  by  this  third  hemorrhage  than  by  either  of  those  preceed- 
ing  it. 

In  some  of  the  principal  points  the  effects  produced  by  the  third 
hemorrhage  corresijonded  very  closely  with  those  of  the  first  and 
seconrl  hemorrhages.  One  of  the  most  imjjortant  factors,  /.  e., 
the  nitrogen  excretion,  showed  the  same  characteristic  of  increas- 
ed elimination.  We  also  see  upon  the  first  day  of  the  experiment 
the  customary  .'^mall  volume  of  urine  having  a  low  specific  grav- 
ity ;  althfjugh  the  difference  between  the  urine  volume  of  this  day 


36 

and  the  volumes  of  the  days  following  was  not  so  great  as  was 
noted  after  the  first  two  hemorrhages.  The  failure  of  the  later 
days  of  this  experiment  to  show  the  usual  percentage  increase  over 
the  volume  of  the  first  day  is  probably  to  be  found  in  the  varia- 
tion in  the  time  of  anaesthesia.  At  each  of  the  first  two  hemor- 
rhages the  beast  was  under  the  influence  of  the  anaesthetic  for 
approximately  2^  hours ;  whereas  at  the  third  hemorrhage  we 
were  forced  to  limit  the  period  of  anaesthesia  to  somewhat  less 
than  i^  hours..  Bearing  in  mind  the  diuretic  eftect  traceable  to 
the  ether  it  is  easily  seen  that  a  much  larger  urine  volume  might 
possibly  have  resulted  had  the  anaesthesia  continued  the  usual 
length  of  time. 

The  length  of  the  period  between  the  operation  and  the  elimi- 
nation of  the  first  urine  appeared  to  differ  very  decidedly  from 
that  of  the  previous  hemorrhages.  At  the  first  and  second  hem- 
orrhages no  urine  was  passed  until  from  16  to  24  hours  after  the 
operation ;  whereas  after  the  third  hemorrhage  the  first  elimina- 
tion was  in  from  5  to  6  hours.  By  an  examination  of  the  data, 
however,  we  learn  that  the  last  urination  preceeded  the  operation 
by  8  hours  and  that  therefore  ys  of  the  normal  daily  urine  volume 
was  probably  in  the  dog's  bladder  at  the  time  of  the  hemorrhage. 
Hence,  under  the  circumstances,  the  passing  of  urine  at  an  earlier 
hour  was  to  have  been  expected.  A  second  urination  occurred 
between  16  and  24  hours  after  the  operation  and  we  think,  there- 
fore, that  we  are  justified  in  considering  the  first  urine  passed  as 
having  been  formed  previous  to  the  operation.  The  volume  pass- 
ed at  this  first  urination  was  180  cc.  which  at  the  same  rate,  would 
have  given  us  a  volume  of  540  cc.  for  the  entire  24  hours.  This 
value  would  agree  very  closely  with  the  average  volume  (530  cc.) 
for  the  experiment  immediately  preceeding. 

The  body  weight  of  the  dog.  fell  from  14.42k.  to  13.76k.  after 
the  hemorrhage,  a  loss  of  0.66k.,  the  greatest  loss  noted  at  any 
time  during  the  84  days  of  the  investigation  (Table  XII). 

2.  Discussion  of  the  Income  and  Outgo  of  Nitrogen,  Sulphur  and 
Phosphorus  After  the  Third  Hemorrhage. — The  third  hemorrhage 
took  place  after  the  animal  had  apparently  entirely  recovered  from 
the  effect  of  the  second  hemorrhage  and  was  again  in  a  condition 
of  approximate  nitrogen  equilibrium.     This  condition  was  repre- 


37 

sen'ted  by  a  loss  of  0.15  gram  of  nitrogen  daily.  Upon  the  day 
of  the  hemorrhage,  as  was  also  the  case  at  each  of  the  preceeding 
hemorrhages,  the  nitrogen  content  of  the  urine  was  the  lowest  of 
any  day  of  the  experiment.  The  second  day,  however,  showed  a 
very  decided  rise  and  the  maximum  was  reached  on  the  third  day. 
The  average  nitrogen  elimination  was  11.669  grams  daily,  where- 
as for  the  five  days  preceeding  the  hemorrhage  the  dog  eliminated 
an  average  of  10.421  grams  daily.  Therefore  the  hemorrhage 
and  its  associated  influences  caused  an  increased  output  of  nitro- 
gen amounting  in  the  aggregate  to  1.248  grams  daily.  It  will  be 
recognized  that  this  was  almost  precisely  the  same  as  the  com- 
bined influence  of  the  anaesthesia,  operation  and  hemorrhage  at 
the  first  withdrawal  of  blood,  the  difl:erence  being  but  0.009 
gram.  The  increase  in  nitrogen  after  the  second  hemorrhage  was 
somewhat  greater  and  was  probably  due,  in  part,  as  has  already 
been  stated,  to  the  cumulative  diuretic  effect  of  the  anaesthesia 
which  appreciably  raised  the  urine  volume.  After  the  third  hem- 
orrhage, however,  as  we  have  seen,  the  anaesthetic  having  been 
administered  for  a  shorter  period,  this  large  urine  volume  was  not 
apparent  but  the  average  volume  for  the  period  was  53  cc.  less 
than  after  the  second  hemorrhage  and  12  cc.  less  than  after  the 
first  hemorrhage  (Table  XII). 

After  making  the  proper  correction  for  the  influence  of  the 
anaesthesia  (See  p.  75)  we  secured  2.185  grams  (Table  XIX) 
as  the  increase  in  the  nitrogen  output  due  to  the  combined  influ- 
ence of  the  operation  and  loss  of  blood  ;  and  correcting  further  for 
the  operation,  we  obtained  as  the  net  influence  of  the  hemorrhage 
an  increase  of  0.479  gram  in  the  daily  nitrogen  elimination.  The 
balance  for  the  last  four  days  of  the  experiment  (Table  IV)  show- 
ed an  increase  of  0.265  gram  in  the  daily  nitrogen  elimination, 
while  the  data  for  the  entire  experiment,  owing  to  the  brevity  of 
the  period  as  well  as  to  the  greatly  increased  elimination  of  the 
first  few  days,  showed  a  daily  increase  of  0.751  gram. 

The  excretion  of  sulphur  apparently  ran  closely  parallel  with 
that  of  nitrogen,  the  sulphur  balance  indicating  a  daily  increase 
of  0.229  gram  (Table  \T).  The  excretion  of  phosphorus  on  the 
other  hanrl.  in  close  agreement  with  the  ob.servations  after  the 
first  anrl  second  hemorrhages,  showed  a  gain  of  0.065  gram  for 
each  day  of  the  period  (Table  VTI). 


38 

Vn.     FOURTH  HEMORRHAGE. 

By  referring  to  the  proper  data  it  will  be  seen  that  the  period 
between  the  third  and  fourth  hemorrhages  was  much  shorter  than 
that  between  the  first  and  second,  or  between  the  second  and  third. 
Furthermore  it  will  be  recognized  that  the  fourth  hemorrhage  was 
instituted  before  the  dog  was  as  near  a  condition  of  nitrogen 
equilibrium  as  upon  the  former  occasions.  It  will  be  remembered 
that  after  the  first  hemorrhage  the  dog  was  brought  to  nitrogen 
equilibrium  in  i6  days,  whereas  21  days  were  necessary  to  secure 
a  like  result  after  the  second  hemorrhage.  Gathering  from  these 
facts  that  it  would  probably  take  three  weeks  or  longer  to  bring 
the  organism  to  nitrogen  equilibrium  after  the  third  hemorrhage, 
we  decided  to  observe  the  influence  of  the  hemorrhage  over  a 
small  number  of  days,  and,  when  we  had  secured  approximate 
equilibrium,  to  follow  with  another  hemorrhage.  Then  as  we  al- 
ready had  data  for  three  hemorrhages  when  the  dog  was  at  nitro- 
gen equilibrium,  we  decided  at  this  point  that  an  interesting  study 
could  be  made  of  hemorrhages  at  short  intervals  upon  this  same 
animal,  no  attempt  being  made  to  secure  nitrogen  equilibrium. 
Therefore  the  period  following  the  third  hemorrhage  was  termi- 
nated, and  the  fourth  hemorrhage  produced,  at  a  point  where  the 
dog  was  losing  0.265  gram  of  nitrogen  daily. 

At  the  time  of  the  fourth  hemorrhage  the  dog  was  able  to  walk 
about  comfortably,  but  was  not  in  as  normal  a  condition  as  upon 
the  occasion  of  the  former  hemorrhages.  The  right  femoral 
artery  just  above  the  point  at  which  the  saphenous  branch  left  it, 
was  selected  as  the  site  of  the  hemorrhage.  After  200  grams  of 
blood  had  been  drawn  the  arterial  pressure  was  so  low  that  only 
a  slight  flow  was  secured.  The  blood  clotted  very  rapidly,  neces- 
sitating frequent  changes  of  the  rubber  tube.  The  body  of  the 
beast  was  freely  massaged  above  the  incision,  and  the  cannula  and 
rubber  tube  were  frequently  cleaned  by  means  of  a  long  platinum 
wire,  but  even  after  using  the  greatest  efforts  to  secure  the  best 
conditions  for  a  satisfactory  hemorrhage  a  slow  dropping  was  the 
maximum  rate  of  flow.  Thinking  that  perhaps  the  clot  might  ex- 
tend back  into  the  artery  the  platinum  wire  was  frequently  insert- 
ed for  some  distance  into  the  vessel  at  the  risk  of  puncture.  After 
securing  about  400  grams  of  blood  the  first  cannula  was  removed 


39 

and  a  second  one  inserted  a  short  distance  above  the  first.  Here 
however  with  fresh  cannula  and  rubber  tube  the  flow  was  not 
rapid  enough  to  prevent  clotting  in  both  tube  and  cannula.  The 
tube  was  discarded  and  blood  collected  directly  from  the  cannula 
but  the  flow  was  not  accelerated.  It  being  evident  that  the  hem- 
orrhage had  continued  to  a  point  where  the  arterial  pressure  was 
insufificient.  the  attempt  to  withdraw  additional  blood  was  aban- 
doned. Therefore  at  10.19  the  administration  of  ether  was  dis- 
continued, the  wound  sewed  up  and  the  animal  returned  to  his 
cage.  The  usual  schedule  follows: — 8.29  A.  M.,  anaesthesia  be- 
gun (ether).  8.37,  trace  of  chloroform.  8.55,  operation  begun. 
9.16.  cannula  inserted.  9.18,  hemorrhage  begun.  9.57,  second  can- 
nula inserted  to  facilitate  flow.  10.18,  hemorrhage  ended,  making 
a  total  collection  of  444.5  grams  of  blood.  10.19,  second  cannula 
out  and  ether  stopped.  Tongue  extremely  w^hite.  10.31,  wound 
sewed  up.  10.36.  in  cage.  10.43,  returning  consciousness.  10.56, 
sits  up.  The  period  of  anaesthesia  of  this  experiment  was  two 
hours  and  fourteen  minutes,  being  somewhat  longer  than  that  of 
the  third  hemorrhage  and  essentially  the  same  as  those  of  the  first 
and  second  hemorrhages.  There  was  practically  no  salivation  at 
this  operation.^  The  total  blood  lost  to  the  organism  was  449.3 
grams. 

I.  Observations  on  the  Influence  of  the  Fourth  Hemorrhage, 
with  a  Discussion  of  the  Income  and  Outgo  of  Nitrogen,  Sulphur 
and  Phosphorus. — An  interesting  condition  was  noted  after  this 
hemorrhage  for  the  first  time  during  our  experiments. 
This  was  a  loss  of  api)etite  on  the  part  of  the  dog. 
At  the  usual  meal  hour  the  animal  was  given  his  food, 
but  contrary  to  custom,  he  ate  very  little  of  the  mix- 
ture, and  a  large  part  of  that  eaten  was  water.  About 
five  minutes  later  he  was  again  offered  the  food  while  he  was 
lying  clown.  This  time  he  ate  the  entire  amount,  but  did  not 
evince  his  accustomed  eagerness.  The  same  lack  of  appetite  was 
noted  upon  the  next  day  also. 

There  was  no  excretion  of  urine  for  12  hours  after  the  hemor- 
rhage (Table  V).     The  urine  volume  on  the  day  of  the  hemor- 

1  Preliminary  to  the  operation  the  wound  from  the  tliird  lit-niorrhage  was  flushed  and 
the  collateral  circulation  noted. 


40 

rhage  was  463  cc.  (Table  XII)  and  this  was  followed  by  a  grad- 
ual rise  until  the  maximum  excretion  of  770  cc.  was  reached  on 
the  third  day.  This  volume  was  also  the  largest  output  of  any 
day  during  the  investigation.  The  usual  decrease  in  body  weight 
was  noticed  after  this  hemorrhage. 

An  examination  of  the  data  showed  a  greatly  increased  nitrogen 
excretion.  This  increase,  after  correcting  for  anaesthesia-opera- 
tion, was  2.506  grams  (Table  XIX)  and  was  about  5  times  as 
great  as  that  observed  after  the  first  and  third  hemorrhages  and 
about  2y2  times  as  great  as  that  after  the  second  hemorrhage. 
This  evidently  showed  quite  forcibly  the  cumulative  efifect  of  suc- 
cessive hemorrhages  upon  proteid  catabolism. 

The  days  following  the  hemorrhages  gave  an  average  increase 
of  0.348  gram  in  the  sulphur  elimination.  In  agreement  with  the 
nitrogen  increase  this  increased  excretion  of  sulphur  was  the 
largest  of  any  period  during  the  investigation. 

For  the  first  time  the  post  hemorrhagic  efifect  upon  the  phos- 
phorus output  was  an  increased  elimination.  There  was  an  aver- 
age daily  increase  of  0.180  gram  (Tables  VII  and  XIX). 

VIII.     FIFTH  HEMORRHAGE. 

No  efforts  were  made  to  get  the  dog  into  nitrogen  equilibrium 
after  the  fourth  hemorrhage,  it  being  thought  best  to  determine 
the  effect  of  another  very  severe  hemorrhage  before  the  organ- 
ism had  been  given  time  to  recover,  in  any  great  degree,  from  the 
loss  of  blood  occasioned  by  the  fourth  hemorrhage. 

The  same  procedure  was  followed  as  at  the  other  hemorrhages, 
the  blood  being  drawn  from  the  right  femoral  artery  above  the 
point  of  the  fourth  hemorrhage. 

X'o  pulse  could  be  felt  at  the  point  determined  upon  for  the  in- 
cision, but  thinking  the  artery  had  changed  position  and  become 
more  deeply  imbedded  in  the  tissues  the  incision  was  made.  After 
locating  the  artery  it  was  found  to  be  completely  closed  by  a  blood 
clot  which  extended  some  distance  above  the  incision  of  the  fourth 
hemorrhage.     This  necessitated  another  incision  farther  up. 

It  was  determined  to  be  guided  entirely  by  the  condition  of  the 
subject  as  to  the  amount  of  the  hemorrhage,  as  the  loss  of  blood 
was  to  be  carried  to  the  extreme  limit. 


41 

After  a  hemorrhage  of  317.5  grams  or  2.46%  of  the  body 
weight  had  been  produced,  it  was  very  apparent,  from  the  condi- 
tion of  the  animal,  that  approaching  death  was  indicated,  and  the 
flow  of  blood  was  suspended.  The  number  of  red  corpuscles  in 
the  blood  at  this  point  was  found  to  be  1,800,000  per  cubic  milli- 
meter, thus  indicating  a  very  anaemic  state. 

In  spite  of  very  vigorous  stirring  in  the  presence  of  25%  NaCl 
solution,  the  blood,  while  possessing  a  very  thin  watery  appear- 
ance as  it  flowed  from  the  artery,  soon  coagulated.  The  whole 
mass  was  filled  with  light  colored  flakes  due  to  the  greatly  in- 
creased number  of  leucocytes. 

After  being  placed  in  his  cage,  and  having  recovered  from  the 
eflfect  of  the  ether,  the  animal  failed  to  show  his  habitual  desire 
to  sit  up,  but  lay  in  a  collapsed  condition  on  the  bottom  of  the 
cage.  At  12  o'clock,  two  hours  after  the  hemorrhage,  his  respira- 
tion was  9  and  his  pulse  152  per  minute.  At  i  P.  M.  152  cc.  of 
urine  was  passed  having  a  specific  gravity  of  1.0255,  rr.uch  high- 
er than  any  former  urine  had  shown  on  the  day  of  hemorrhage. 
At  4  P.  M.  his  extremities  were  cold,  axillary  temperature  was 
subnormal,  and  in  spite  of  the  greatest  efforts  to  preserve  his  life 
the  animal  died  3  hours  later. 

I.  Post-mortem  Examination. — The  bladder  was  removed  and 
14  cc.  of  urine  with  a  specific  gravity  of  1.036  (the  highest  of  any 
sample  during  the  entire  investigation)  was  secured.  From  our 
records  it  was  learned  that  the  dog  urinated  last  at  i  P.  M.  and 
therefore  this  14  cc.  of  urine  represented  the  total  urine  formation 
for  f}  hours.  Ordinarily  the  animal  would  have  passed  over  100 
cc.  during  a  period  of  that  length. 

All  of  the  animal's  organs  were  extremely  pale,  and  showed 
practically  no  tendency  to  bleed  when  cut.  Even  the  heart  was 
blanched  and  upon  laying  it  open,  a  very  small  clot  in  the  left 
ventricle  was  the  only  blood  to  be  found. 

In  general  the  i)Ost-mortem  examination  showed  conclusively 
that  a  very  great  portion  of  the  animal's  blood  had  been  removed. 
As  only  about  300  cc.  of  blood  was  taken  at  the  fifth  hemorrhage, 
it  was  evident  that  the  regeneration  of  the  volume  during  the 
period  since  the  fourth  hemorrhage  had  been  exceedingly  slow. 


42 


C— SECOND  SERIES  OF  EXPERIMENTS. 


This  second  series  of  experiments  was  much  shorter  than  the 
first  series  and  was  made  principally  for  the  purpose  of  check- 
ing, upon  another  organism.,  the  results  from  our  first  series.  In 
this  second  series  the  experiments  were  made  upon  a  dog  weigh- 
ing 11.85kg. 

In  our  first  series  the  influence  of  anaesthesia-operation  was  not 
determined  until  after  the  animal  had  been  subjected  to  a  prelim- 
inary hemorrhage.  In  the  present  series,  however,  after  getting 
the  organism  into  a  condition  of  approximate  nitrogen  equilibrium 
by  means  of  a  preliminary  period,  the  influence,  of  anaesthesia- 
operation  was  studied  before  any  blood  was  drawn.  Following 
this,  after  returning  to  equilibrium,  the  efifect  of  hemorrhage  was 
studied. 

The  diet  as  shown  by  Table  XV,  was  qualitatively  the  same  as 
that  of  the  first  series  of  experiments. 

I.     Preliminary  Experiment. 

In  a  preliminary  period  of  nine  days,  during  which  the  dog  was 
fed  a  constant  diet,  the  organism  was  brought  to  approximate 
nitrogen  equilibrium.  For  this  period,  as  may  be  seen  from  an 
examination  of  Table  XVI,  there  was  an  average  daily  loss  of 
but  o.io  gram  of  nitrogen  to  the  body.  The  organism  was  there- 
fore in  an  exceptionally  good  condition  for  the  study  of  the  influ- 
ence of  anaesthesia-operation. 

II.     Influence  of  Anaesthesia-Operation. 

As  has  been  said,  the  influence  of  anaesthesia-operation  in  the 
first  series  of  experiments  was  not  determined  until  after  the  or- 
ganism had  been  subjected  to  an  initial  hemorrhage.  In  order  to 
secure  additional  data  the  influence  of  anaesthesia-operation  was 
determined  in  the  second  series  before  the  first  hemorrhage. 

As  usual,  the  femoral  artery  was  laid  bare  just  above  the  point 
at  which  the  saphenous  branch  is  given  off.  The  incision  was 
longer  and  deeper  than  at  any  previous  operation,  in  order  to 
demonstrate  to  the  full  any  influence  exerted  by  this  operative 


43 

procedure.  The  schedule  of  operations  was  as  follows : — 8.22  A. 
M..  anaesthesia  begun.  8.26,  trace  of  chloroform  given.  8.55, 
incision  made.  9.20,  wound  sewed  up.  9.45,  returning  conscious- 
ness. 10.00.  dog  sat  up.  This  animal  was  evidently  more  suc- 
ceptible  to  the  influence  of  the  ether  than  the  former  beast  as  was 
evidenced  by  the  ease  with  which  the  preliminary  anaesthesia  was 
conducted  as  well  as  by  the  weakness  of  the  animal  after  con- 
sciousness had  returned. 

The  influence  of  anaesthesia-operation  was  studied  through  a 
period  of  nine  days.  Here  as  in  the  case  of  the  first  series  of  ex- 
periments, the  anaesthesia-operation  was  followed  by  an  imme- 
diate rise  in  the  excretion  of  nitrogen  by  the  urine.  The  average 
daily  output  of  nitrogen  by  the  urine,  during  the  preliminary 
period  was  'j.'/'ii  grams  (Table  XVII).  On  the  first  day  of  the 
anaesthesia-operation  period  the  nitrogen  elimination  was  increas- 
ed to  8.56  grams  (Table  X\TI)  and  the  average  daily  output  for 
the  first  three  days  of  the  period  was  9.12  grams.  Taking  the 
whole  nine  days  into  consideration  the  average  daily  output  of 
nitrogen  by  the  urine  was  8.79  grams.  The  balance  for  the  period 
showed  a  daily  loss  of  1.09  grams  of  nitrogen  to  the  organism 
(Table  X\'I).  At  the  beginning  of  the  period  the  organism  was 
losing  o.io  gram  of  nitrogen  daily,  and  correcting  for  this  we  see 
that  the  net  effect  of  the  anaesthesia-operation  upon  the  excretion 
of  nitrogen  has  been  to  increase  the  output  on  an  average  0.99 
gram  per  day. 

The  general  tendency  of  the  sulphur  excretion  to  follow  a 
course  similar  to  that  of  the  nitrogen  excretion  was  noted  here. 
By  means  of  the  preliminary  period  of  nine  days  the  organism 
was  placed  in  a  condition  of  almost  exact  sulphur  equilibrium,  the 
average  daily  loss  being  0.00 1  gram  (Table  XVIII).  The  in- 
fluence of  the  anaesthesia-operation  was  shown  in  the  form  of  an 
increased  sulphur  excretion.  During  this  period  the  organism 
showed  an  average  daily  loss  of  0.053  gram,  thus  signifying  that 
the  sulphur  excretion  was  increased  0.052  gram  daily  as  the  ef- 
fect of  the  anaesthesia-operation,  or  a  total  increase  of  0.496  gram 
for  the  nine  days. 

As  usual  the  phosphorus  excretion  showed  a  slight  decrease  as 
the  effect  of  the  anaesthesia-operation.     The  animal  at  the  open- 


44 

ing  of  the  period  was  losing  0.03  gram  of  phosphorus  daily 
(Table  XVIII).  Under  the  intiuence  of  the  anaesthesia-operation 
the  phosphorus  excretion  decreased  a  trifle,  making  the  average 
daily  loss  for  the  period  0.025  gram. 

III.     Influence  of  Hemorrhage. 

Upon  June  20  a  hemorrhage  was  induced  in  the  left  femoral 
artery.  The  blood  was  withdrawn  rapidly  in  this  case,  the  total 
amount  of  blood  taken  aggregating  342.5  grams  or  3.11%  of  the 
body  weight  of  the  animal.  Due  to  the  rapidity  of  the  hemor- 
rhage there  were  profound  disturbances  of  pulse  and  respiration, 
but  these  were  of  short  duration.  The  schedule  of  operations  was 
as  follows: — 8.15  A.  M.,  anaesthesia  begun.  8.20,  chloroform 
given.  8.35,  operation  begun.  8.50,  cannula  inserted.  8.53, 
hemorrhage  begun.  9.00,  hemorrhage  ended.  9.02,  cannula  re- 
moved. 9.15,  wound  sewed  up.  9.20,  anaesthesia  discontinued 
and  dog  returned  to  cage.    9.50,  dog  stood  up. 

In  complete  agreement  with  the  data  for  the  combined  effect  of 
hemorrhage  and  anaesthesia-operation  in  the  first  series  of  exper- 
iments, we  also  find  here  in  the  second  series  that  a  considerable 
increase  in  the  excretion  of  nitrogen  by  the  urine  resulted.  The 
average  daily  excretion  of  nitrogen  was  9.47  grams  for  the  period 
of  five  days  following  the  hemorrhage  (Table  XVII).  The 
nitrogen  balance  for  this  period  showed  an  average  daily  loss  of 
1.88  grams  (Table  XVI).  Correcting  this  for  the  normal  daily 
loss  of  o.io  gram,  determined  in  the  preliminary  period,  we  find 
that  the  combined  effect  of  hemorrhage  and  anaesthesia-operation 
has  been  to  cause  an  average  daily  increase  of  1.78  grams  in  the 
excretion  of  nitrogen.  And  correcting  further  for  the  influence 
of  anaesthesia-operation  (0.99  gram),  we  find  that  the  withdrawal 
of  342.5  grams  of  blood^  was  instrumental  in  causing  an  average 
daily  increase  of  0.79  gram  in  the  nitrogen  excretion  throughout 
a  period  of  five  days. 

As  usual  the  course  of  the  sulphur  excretion  was  similar  to 
that  of  nitrogen,  while  the  phosphorus  excretion  instead  of  being 
increased  by  the  hemorrhage  was  slightly  decreased  (Table 
XVIII). 

IThe  analysis  of  this  blood  was  as  follows  :  Nitrogen,  3.17  per  cent.;  sulphur,  0.156  per 
cent.;  phosphorus,  0.48  per  cent. 


45 


D.— ALTERATIONS  IN  THE  SPECIFIC  GRAV- 
ITY, AND  IN  THE  NITROGEN,  SULPHUR 
AND  PHOSPHORUS  CONTENT  OF  THE 
BLOOD  FOLLOWING  HEMORRHAGE. 

The  first  and  second  hemorrhages  being  separated  from  each 
other  by  a  period  of  nearly  six  weeks  may  be  considered  initial 
hemorrhages.  That  is,  the  composition  of  the  blood  drawn  on 
Dec.  23  cannot  be  considered  as  the  composition  of  the  blood  as 
effected  by  the  hemorrhage  of  Nov.  14,  for  it  is  well  known  that 
the  influence  of  a  very  severe  hemorrhage  would,  so  far  as  the 
composition  of  the  blood  is  concerned,  have  disappeared  long  be- 
fore the  second  hemorrhage.  Hence  the  blood  drawn  on  Dec.  23 
may  be  considered  normal  blood. 

Of  the  elements  under  consideration  the  nitrogen  appeared  to 
be  the  only  one  effected  in  any  uniform  manner  by  the  successive 
hemorrhages.  With  this  element  beginning  Dec.  23,  which  may 
be  considered  the  commencement  of  the  series  of  hemorrhages, 
we  note,  by  referring  to  Table  II,  the  normal  nitrogen  content  of 
the  blood  to  be  2.85%.  Three  weeks  later  the  third  hemorrhage 
showed  a  nitrogen  content  of  2.38^,  while  following  this,  after 
an  interval  of  one  week,  the  fourth  hemorrhage  gave  us  a  nitrogen 
value  of  1.848%,  and  after  a  short  interval  of  four  days  at  the 
fifth  and  final  hemorrhage  the  minimum  point  v/as  reached  and 
the  surprisingly  low  nitrogen  value  of  1.421%  was  recorded.  Thus 
the  effect  of  four  successive  hemorrhages  in  a  period  of  one 
month  on  the  nitrogen  content  of  the  blood  had  been  to  decrease 
the  amount  from  2.85%  to  1.421%,  a  decrease  of  more  than  50%. 
There  was  an  actual  decrease  of  16%;  in  the  nitrogen  content  of 
the  blood  at  the  third  hemorrhage,  of  22%  at  the  fourth  hemor- 
rhage and  of  23%  at  the  fifth  hemorrhage.  The  time  between  the 
hemorrhages  decreasing  as  it  did  from  three  weeks  to  only  four 
days  at  the  final  hemorrhage,  there  seems  to  have  been  a  very  pro- 
nounced cumulative  effect  of  some  kind  which  produced  in  four 
days  time  a  greater  percentage  decrease  in  the  content  of  nitrogen 
than  was  previously  secured  after  a  lapse  of  three  weeks. 


46 

The  sulphur  did  not  exhibit  the  same  regularity  as  the  nitrogen. 
It  coincided,  however,  in  showing  the  minimum  output  following 
the  last  hemorrhage.  The  content  of  phosphorus  also  was  not 
governed  by  any  uniformity,  but  differed  radically  from  the  nitro- 
gen and  sulphur  in  showing  the  maximum  percentage  at  the  last 
hemorrhage.  Hence  the  maximum  phosphorus  occurred  coin-, 
cidently  with  the  minimum  sulphur,  and  the  minimum  phosphorus 
with  the  maximum  sulphur  (See  analyses  for  fourth  hemor- 
rhage). 

The  specific  gravity  was  lowered  at  each  successive  hemor- 
rhage, falling  from  1.0625  ^^  ^^^  ^^^^  hemorrhage  to  1.047  ^^  the 
fourth.  The  blood  from  the  fifth  hemorrhage  showed  a  greatly 
increased  specific  gravity.  This  was  due  to  the  fact  that  the  blood 
immediately  after  withdrawal,  even  in  the  presence  of  the  25% 
NaCl  solution,  showed  a  tendency  to  coagulate.  This  of  course 
prevented  the  accurate  determination  of  its  specific  gravity,  and 
made  the  specific  gravity  as  determined  much  too  high.  We  are 
satisfied  that  the  blood  as  drawn  from  the  animal  had  a  very  low 
specific  gravity  as  it  was  extremely  thin  and  watery.  Without 
doubt  it  was  lower  than  that  of  any  of  the  samples  previously  ex- 
amined. 


E.— RELATION  BETWEEN  TOTAL  NITROGEN 
AND  VOLUME  OF  URINR 


The  ratio  between  the  nitrogen  content  of  the  urine  and  the 
urine  volume  showed  quite  decided  fluctuations  under  different 
experimental  conditions.  Starting  with  a  ratio  of  i  :55-9  for  the 
preliminary  experiment  (Table  XIII),  the  first  hemorrhage  pro- 
duced a  very  decided  lowering  of  the  ratio,  the  relation  at  this 
time  being  i  145 .8.  During  the  latter  part  of  the  first  hemor- 
rhage experiment  the  ratio  gradually  rose  and  at  the  time  nitro- 
gen equilibrium  had  been  reached  it  was  i  :54.5. 

The  anaesthetic  through  its  diuretic  action  now  forced  the  ratio 
still  higher,  the  maximum  point  being  reached  during  the  last  days 
of  the  anaesthesia  experiment.  This  was  evidently  due  to  the 
cumulative  diuretic  effect  of  the  ether.     From  the  result  obtained 


47 

during  the  first  days  of  the  anaesthesia  experiment  we  would  natu- 
rally expect  a  rise  in  the  ratio  during  the  first  days  of  the  anaes- 
thesia-operation experiment.  Here,  however,  although  the  anaes- 
thesia was  of  the  same  duration  as  before,  instead  of  a  rise  we  ob- 
tained a  fall.  This  evidently  indicated  that  the  individual  influ- 
ence of  the  operation  had  been  partly  to  overcome  the  rise  due  to 
anaesthesia.  The  ratio  rose  slightly  higher  during  the  following 
days  of  the  experiment,  but  was  at  no  time  higher  than  the  normal 
ratio  of  the  preliminary  experiment. 

The  second  hemorrhage  produced  a  corresponding  fall  to  that 
observed  after  the  first  hemorrhage  with  a  similar  rise  dtiring  the 
later  days  of  the  experiment.  The  third,  fourth  and  fifth  hemor- 
rhages, while  not  strictly  comparable  with  the  first  and  second 
hemorrhages,  each  showed  a  low  ratio,  those  of  the  fourth  and 
fifth  (I  143.0  and  i  140.3)  being  the  lowest  of  the  whole  investiga- 
tion. This  rapid  fall  in  the  ratio  after  the  fourth  and  fifth  hemor- 
rhages was  evidently  due  to  the  fact  that  while  the  regenerative 
activities  caused  a  large  increase  in  proteid  decomposition  a  cor- 
responding increase  in  water  output  was  impossible  since  the 
water  content  of  the  body  had  been  so  greatly  decreased  by  the 
excessive  blood-letting. 


R— DISCUSSION  OF  RESULTS. 


The  data  from  our  experiments  indicate  that  the  loss  of  blood 
by  an  organism  is  followed  immediately  by  an  increased  proteid 
catabolism.  And  furthermore,  it  was  plainly  demonstrated,  by  a 
series  of  hemorrhages  on  the  same  organism  brought  to  a  condi- 
tion of  nitrogen  equilibrium  before  each  loss  of  blood,  that  there 
is  a  tendency  under  such  conditions  for  the  catabolism  of  proteid 
to  become  more  and  more  pronounced  as  the  series  progresses. 
This  was  shown  by  a  daily  increase  of  1.257  grams  in  the  nitrogen 
elimination  as  the  result  of  a  hemorrhage  of  2.93%  of  the  body 
weight  of  the  animal,  followed  some  weeks  later  by  a  daily  in- 
crease of  3.275  grams  in  the  nitrogen  elimination  after  a  hemor- 
rhage of  3.26%  of  the  body  weight. 


48 

The  influence  of  the  anaesthesia  to  which  the  animal  was  sub- 
jected was  to  cause  a  daily  decrease  of  0.937  gram  in  the  nitrogen 
output.^  Another  influencing  factor  was  the  operation  which  was 
shown  of  itself  to  cause  an  increased  proteid  catabolism,  the  aver- 
age daily  nitrogen  elimination  being  increased  1.706  grams.  Now 
taking  the  values  named  in  the  above  paragraph  as  representing 
the  gross  influence  of  the  hemorrhage,  and  correcting  them  for 
the  influence  of  the  anaesthesia  and  of  the  operation,  we  see  that 
the  net  efl^ect  of  the  hemorrhage  alone  (Table  XIX)  was  to  cause 
a  daily  increase  in  the  nitrogen  elimination  varying  from  0.488 
gram  after  a  loss  of  blood  amounting  to  2.93%  of  the  body  weight, 
to  2.506  grams  after  a  hemorrhage  of  3.26%  of  the  body  weight. 

The  influence  of  the  hemorrhage  and  its  accompanying  factors 
upon  the  excretion  of  sulphur  was  similar  to  that  noted  in  the  case 
of  nitrogen.  The  daily  increase  in  the  sulphur  excretion  follow- 
ing hemorrhage  varied  from  0.023  gram  to  0.348  gram,  and  the 
increase  due  to  the  anaesthesia-operation  was  0.119  gram  daily. 
The  anaesthesia  of  itself  caused  a  daily  decrease  of  0.128  gram 
in  the  sulphur  excretion. 

The  phosphorus  excretion,  with  a  single  exception,  was  influ- 
enced by  hemorrhage  and  its  associated  factors  in  a  manner  di- 
rectly opposite  to  that  in  which  the  nitrogen  and  sulphur  excre- 
tions were  influenced.  In  the  case  of  the  phosphorus  the  net  in- 
fluence of  the  hemorrhage  alone  was  to  cause  a  decrease  in  the 
elimination,  the  daily  decrease  varying  from  0.163  gram  to  0.065 
gram.  The  anaesthesia-operation  caused  a  daily  decrease  of  0.148 
gram  and  the  anaesthesia  alone  caused  a  daily  increase  of  0.108 
gram.  The  decrease  in  the  elimination  of  phosphorus  after  hem- 
orrhage we  believe  to  be  due  primarily  to  the  unusual  demands 
made  upon  nuclear  material  for  the  construction  of  leucocytes  and 
nucleated  erythrocytes. 

Upon  the  day  of  the  hemorrhage  the  urine  volume  was  invar- 
iably sub-normal,  but  following  this  came  a  greatly  increased  out- 
put of  urine  which  generally  reached  its  maximum  on  the  third 
or  fourth  day  after  the  hemorrhage.  This  increase  in  the  urine 
volume  was  partly  due  no  doubt  to  the  diuretic  influence  of  the 
anaesthetic.     Furthermore,  the  animal  received  a  uniform  supply 

IHawk  :     Proceedings  of  the  American  Physiological  Society,  1903. 


49 

of  water  daily  and  this  supply  was  sufficient  for  the  dog  under 
normal  conditions  as  was  witnessed  by  the  promptness  with  which 
nitrogen  equilibrium  was  reached.  Upon  withdrawing  a  large 
amount  of  blood,  however,  this  normal  water  supply  was  more 
than  sufficient  for  the  needs  of  the  altered  metabolism,  and  there- 
fore a  portion  of  the  excess  was  excreted.  As  was  to  have  been 
expected  this  large  urine  volume  was  accompanied  by  an  increased 
elimination  of  nitrogen,  this  increase  being  due  in  part  to  a  flush- 
ing-out of  the  tissues  as  well  as  to  a  direct  stimulation  of  proteid 
catabolism.^ 

The  body  weight  of  the  animal  decreased  very  materially  on  the 
day  of  the  hemorrhage,  the  decrease  being  due  apparently  as  much 
to  the  influence  of  the  anaesthesia-  as  to  the  effect  of  the  loss  of 
blood. 

The  well  konwn  leucocytosis  followed  the  hemorrhages  and  a 
decrease  in  the  number  of  erythrocytes  was  also  noted.  As  has 
already  been  stated  on  page  41,  a  most  remarkable  leucocytosis 
was  observed  at  the  time  of  the  fifth  hemorrhage. 

The  appetite  of  the  subjects,  with  the  single  exception  noted, 
(page  39).  continued  uniformly  good  throughout  the  experi- 
ments, even  when  the  body  weight  of  the  animal  had  decreased 
very  considerably.  The  regular  normal  diet  which  was  just  suffi- 
cient to  bring  the  organism  weighing  about  17kg.  to  nitrogen 
equilibrium  was  very  probably  in  excess  of  the  needs  of  the  body 
when  the  weight  had  been  reduced  to  about  13kg.  at  the  time  of 
the  fourth  hemorrhage.  Therefore  this  excessive  amount  of  pro- 
teid material  may  have  tended  to  stimulate  proteid  catabolism  and 
thus  assisted  in  the  production  of  the  maximum  increase  in  the 
nitrogen  elimination  which  was  observed  at  that  time. 


G.— CONCLUSIONS. 


I.  Hemorrhages  of  2.9%  to  3.5%  of  the  body  weight  u])on 
dc^s  fed  a  constant  diet  and  in  a  condition  of  nitrogen  equilib- 
rium, caused  an  increased  urinary  elimination  of  nitrogen  and  sul- 
phur and  a  decreased  elimination  of  phosphorus. 

'  Hawk  :     fniversity  of  Pennsylvania  Medical  Bulletin,  igo.s.XVIII.,  7. 
2  Hawk  :     Proceedings  of  the  American  Pliysiological  Society,  1903. 


50 

2.  Ether  anaesthesia  incident  to  ordinary  surgical  procedure 
was  followed  by  a  decreased  elimination  of  nitrogen  and  sulphur 
and  an  increased  elimination  of  phosphorus  and  chlorine. 

3.  The  combined  effect  of  the  ether  anaesthesia  and  the  opera- 
tive procedure  was  to  cause  an  increased  elimination  of  nitrogen, 
sulphur  and  chlorine  and  a  decreased  elimination  of  phosphorus. 

4.  After  hemorrhages  of  2.9%  to  3.5%  of  the  body  weight  there 
was  an  immediate  decrease  in  the  volume  of  urine.  This  decrease 
was  followed  after  the  first  day  by  an  increase  and  the  maximum 
urine  volume  generally  occurred  on  the  third  or  fourth  day  after 
the  hemorrhage. 

5.  Ether  anaesthesia  unaccompanied  by  loss  of  blood,  produced 
an  immediate  diuresis  which  caused  the  maximum  urine  volume 
to  appear  vipon  the  day  of  the  anaesthesia. 

6.  Successive  hemorrhages  on  the  same  organism  caused  a 
gradual  decrease  in  the  nitrogen  and  sulphur  content  of  the  blood, 
and  a  less  regular  decrease  in  the  phosphorus  content. 

7.  In  a  series  of  hemorrhages  the  specific  gravity  of  the  blood 
gradually  decreased  to  the  end  of  the  series. 

8.  Repeated  hemorrhages  at  intervals  of  from  one  to  five  weeks 
caused  the  coagulation  rate  of  the  blood  to  shorten  very  percepti- 
bly, particularly  at  the  end  of  the  series. 

9.  A  decrease  in  body  weight  followed  hemorrhage. 

10.  There  was  a  pronounced  leucocytosis  and  a  decrease  in  the 
number  of  erythrocytes  after  hemorrhage. 

11.  Hemorrhage  caused  no  disturbance  of  the  digestive  func- 
tions and  had  no  effect  upon  intestinal  putrefaction. 

The  author  wishes  to  express  his  sincere  thanks  to  Prof.  W.  J. 
Gies  under  whose  direction  this  investigation  was  conducted. 


H.— BIBLIOGRAPHY, 


1.  Bauer  :     Zeitschrift  fiir  Biologie,  1872,  VIII.,  p.  567. 

Miinchener  Medicinische  Wochenschrift,  1892,  39,  p.  537 

2.  Jiirgensen  :     v.  Ziemssen's  Handbuch  d.  Allgenieine  Therapie,   1880. 

3.  Maltschewsky  :     Thesis  (Russian),  1892. 

4.  Ascoli  and  Draghi  :     Berliner  Klinische  "Wochenschrift,   1900,   37,  p. 
1055- 


51 

5-     Frankel  :     Virchow's  Archiv,  1876,  LXVII. 

6.  V.  Noorden  :     Lehrbuchder  Pathologic desStofFwechsels,  Berlin,  1893. 

7.  Lukjanow  :     Zeitschrift  fiir  Phys.  Chem.  1883,  VIII.,  p.  336. 

8.  Giirber  :     Miinchener  ]Medicinische  Wochenschrift,   1892,  39,  pp.  416 
and  605. 

9.  Autokonenko  :     Archives  des  sciences  biologiques,    1893,   II.,  p.  516. 

10.  Baumann  :    Journal  of  Physiology,  1903,  29,  p.  18. 

11.  Luzet  :     Archives  de  physiologic  nortnale  et  pathologiquc,  1891,  3, 
P-  455- 

12.  Vulpain  :     Comptes  rendus  des  sceances  de  I'acadcmic  des  sciences, 
1877,  84,  p.  1279. 

13.  Hiinerfauth  :     Virchow's  Archiv,  1879,  76,  p.  310. 

14.  Buntzen  :     Copenhagen,  1879,  p.  56.     Quoted  by  Jiirgensen. 

15.  V.  Lesser:     Archiv  fiir  Anatomic  und   Physiologic  ( Physiologische 
Abtheilung),  1S78,  p.  74, 

16.  Lyon  :     Virchow's  Archiv,  84.     Quoted  by  Autokonenko. 

17.  Bizzozcro  and  Salvioli  :     Arch,  per  lescienze  mediche,  IV.,  12,  p.  273. 

18.  Monassein  :     Tiibingen,  1872,  p.  41. 

19.  Tschoudnowsky  :     Thesis  (Russian),  1869.     Quoted  by  Autokonenko. 

20.  Riedcr  :     Bcitriige  zur  Kenntniss  der  Lencocytosis  und  verwandter 
zustande  des  Blutes,  1892,  p.  90. 

21.  Haycm  :     Lecons  sur  les  modifications  du  sang,  Paris,  1882. 

22.  Himmelstjerna  :     Dissertation,  1882,  p.  25.  Quoted  by  Autokonenko. 

23.  Virchow  :     Die  Cellularpathologie,  Berlin,  1862. 

24.  Limbeck  :     Grundriss  einen  clinischcn  pathologic  des   Blutes,  1892, 
p.  49- 

25.  Molassez  :     Gazette  Medicale  de  Paris,   1880,  II.,  p.  465. 

26.  Henle  :     Quoted  by  Jiirgensen  and  Autokonenko. 

27.  Rcmak  :     Quoted  by  Jiirgensen  and  Autokonenko. 

28.  Woltersom  :     Quoted  by  Donders,   Phys.   des  Menschen,  I.,  p.  166. 
Quoted  by  Bauer. 

29.  Conheim  :  ] 

30.  Massart  and  Bordet  :  I 

31.  Gabritschewsky  :  j 

32.  Afanasiew  :  I 

33.  Leber  :  j-  Quoted  by  Autokonenko. 

34.  Roemer  :  I 

35.  Buchner  : 

36.  Escherich  : 

37.  Foster :  J  .  ' 

38.  Leichtenstcrn  :     Zeitschrift  fiir  Biologic,  1868,  VII.,  p.  215. 

39.  Kussniau!   and  Tenner  :     Frankfort,    1857.     Quoted   by    Bauer   and 
Jiirgen.sen. 

40.  Naunyn  and  Quinke  :     Du  Bois  Reymond's  Archiv,  1869,  Heft  2. 

41.  Gies  :     American  Journal  of  Physiology,  1903,  IX.,  p.  13. 


52 

42.  Worm-MuUer  :     Berichte  der  Sachsischen  Gesellschaft  der  Wissen- 
schaften,  1873,  p.  642. 

43.  Volkmann  :     Die  Hamodynamik,  p.  197. 


44 
45 
46 

47 
48 

49 
50 
51 
52 
53 
54 
55 
56 
57 
58 

59 
60 
61 
62 

63 
64 

65 

66 

67 
von 
68 
69 
70 
71 
72 
73 
74 


Nawrotzky  : 

Magendie  :  [Quoted  by  Bauer. 

Nawlichen  : 

Hayem  :     Loc.  cit. 

Hall  :     Quoted  by  Liebernieister  in  v.  Ziemmsen's  Handbuch,  1880, 1. 

Traube  :     Quoted  by  Bauer. 

Spielnian  :  1 

Frese  and  Charaszewski  :       >  Quoted  by  Jiirgensen. 

Wunderlich  :  J 

Briicke  :     Virchow's  Archiv,  1857,  XII.,  p.  179. 

Nasse  :     Das  Blut,  Bonn,  1836,  p.  156. 

Jiirgensen  :     Loc.  cit. 

Mayer  :     Quoted  by  Jiirgensen. 

Vierordt  :     Archiv  der  Heilkunde,  XIX.,  p.  193. 

Briicke  :     Loc.  cit. 

Nasse  :     Loc.  cit. 

Baumann  :     Loc.  cit. 

Sherrington  and  Copenian  :     Journal  of  Physiology,  1893,  XIV  ,  p.  52. 

[■Quoted  by  Sherrington  and  Copeman. 


Jones : 

Bizzozero  and  Salvioli  :     Loc.  cit. 
Woltersoni  :     Loc.  cit.     Quoted  by  Vierordt. 
Baumann  :     Loc.  cit. 

Tolmatscheff :     Medicinische  Chem.  Untersuchungen  herausgegeben 
Hoppe-Seyler,  Tiibingen,  III.,  p.  400. 
Bauer  :     Loc.  cit. 
Jiirgensen  :     Loc.  cit. 
Lister  :     Quoted  by  Jiirgensen. 
Hiinerfauth  :     Loc.  cit. 

Goll  :     Zeitschrift  fiir  Rationelle  Medicin,  Neue  Folge,  1854,  IV.,  p.  78. 
Schramm  :     Medic.  Jahrbiicher,  Wien,  1885,  p.  492. 
V.  KireefF:     Archiv  fiir  Anatomic  und  Physiologic   (Physiologische 
Abtheilung),  1884,  p.  156. 

75.  Hayem  :     Loc.  cit. 

76.  Noll  :     Quoted  by  Kronecker. 

77.  Kronecker  :     Correspondenzblatt  fiir  Schweizer  Aerzte,  1886,  16. 

78.  Ott :     Thesis  (Russian),  1884,  p.  32. 

79.  Maydl  :     Medic.  Jahrbiicher,  Wien,  1884,  p.  77. 

80.  Schramm  :     Loc.  cit. 

81.  Prevost  and  Dumas  :     Quoted  by  Maydl. 

82.  Skvortsov  :  Inaugural  Dissertation,  St.  Petersburg,  1890.  Quoted 
in  Bulletin  45,  Office  of  Experiment  Stations,  Department  of  Agriculture* 
U.  S.  A. 


53 

S3.     Debierre  and  Linossier:     Bulletin  General  de  Therapeutique,  1885, 
CVIII,  p.  167. 

84.  Emminghaus  :     Bar.  uber  d.  Verhandl.  d.  Kouig. 

Sachs  ;  Gesellsch.   d.  Wiss.   Z.  Leipzig,  1S73,  XXV. 

85.  Fleischer  and  Penzoldt  :     Deutsches  Archiv  fiir  Klinische  Medicin, 
1880,  XXVI.,  p.  400. 

86.  Lipmann-Wulf  :     v.  Noorden's  Beitrage  zur  Lehre  vom  Stoffwechsel, 
1892,  I.,  24. 

87.  Ketcher  :     Vrach,  11,  p.  1042.     Quoted  in  Bulletin  45  (loc.  cit). 

88.  Moraczewski  :     Zeitschrift  fiir  Klinische  Medicin,  1897,  33. 

89.  Eichorst  :     Die   progress,    pernic.     Anamie,    Leipzig,    1S78,    p.    205. 
Quoted  by  v.  Noordeu. 

90.  Burzhinski  :     Vrach,  10,  p.  994.     Quoted  in  Bulletin  45,  (loc.  cit). 

91.  Sticker  :     Zeitschrift  fiir  Klinische  Medicin,  1888,  14. 

92.  Albrecht  :     Jahrbuch  fiir  Kinderheilkunde,  1882,  18,  p.  i. 

93.  Hayem  :     Loc.  cit. 

94.  Rieder  :     Loc.  cit. 

95.  Hayem  :     Loc.  cit. 

96.  Samuel  :     Manuel  de  pathologic  general,  1S79,  p.  254. 

97.  King  :     American  Journal  of  the  Medical  Sciences,  1902,  CXXIV., 
p.  450. 

98.  Mosler  :     Leukamie.     Quoted  by  Bauer. 

99.  Lowit  :     Virchow's  Archiv,  117,  p.  569. 

100.  Miiller  :     Deutches  Archiv  fiir  Klinische  Medicin,  1901,  48. 
loi.     Osier:     Osier's  Practice  of  Medicine,  p.  789. 

102.  V.  Noorden  :     Loc.  cit. 

103.  Neusser  :     Wiener    Klinische    Wochenschrift,     1897,     10,    p.    629. 
Quoted  by  Kolisch. 

104.  Kolisch  :     Wiener  Klinische  Wochenschrift,  1897,  26,  p.  628. 


I.— BIOGRAPHICAL. 


Philip  Bovier  Hawk  graduated  from  Wesleyan  University  in 
1898  with  the  degree  of  Bachelor  of  Science.  During  the  next 
two  years  (1898-1900)  he  was  assistant  to  Prof.  W.  O.  Atwater 
of  We.sleyan  University.  During  this  time  he  also  did  graduate 
work  in  the  University  and  received  the  degree  of  Master  of 
Science  in  1900.  As  a  university  scholar,  in  1900- 1901,  he  pur- 
sued graduate  studies  in  physiological  cheniistr}^  and  physiology 
in  Yale  University  and  in  1902  received  the  degree  of  Master  of 
Science  from  that  institution.     He  served  as  assistant  in  physi- 


54 

ological  chemistry  in  Columbia  University  (College  of  Physicians 
and  Surgeons)  during  1901-1903,  at  the  same  time  pursuing 
graduate  work  in  physiological  chemistry.  In  1903  the  degree 
of  Doctor  of  Philosophy  was  conferred  upon  him  by  Columbia 
University. 

He  is  a  member  of  Delta  Kappa  Epsilon,  Sigma  Xi,  the  Ameri- 
can Physiological  Society  and  the  Society  for  Experimental 
Biology  and  Medicine. 


J.— PUBLICATIONS. 


1.  On  the  elimination  of  nitrogen,  sulphates  and  phosphates  after  the 
ingestion  of  proteid food  (with  H.  C.  Sherman);  American  Journal  of  Physi- 
ology, 1900,  IV.,  p.  25. 

2.  Chemical  studies  of  osseomucoid,  with  determinations  of  the  heat  of 
combustion  of  some  connective  tissue  glucoproteids  (with  W.  J.  Gies); 
American  Journal  of  Physiology,  1901,  V.,  p.  387. 

3.  On  the  composition  and  chemical  properties  of  osseoalbumoid,  with  a 
comparative  study  of  the  albumoid  of  cartilage  (with  W.  J.  Gies);  American 
Journal  of  Physiology,  1902,  VII.,  p.  340. 

4.  On  the  quantitative  determination  of  acid  albumin  in  digestive  mix- 
tures (with  W.  J.  Gies);  American  Journal  of  Physiology,  1902,  VII.,  p.  460. 

5.  On  the  influence  of  rennin  upon  the  digestion  of  the  proteid  constitu- 
ents of  milk  ;  American  Journal  of  Physiology,  1903,  X.,  p.  37. 

6.  On  the  time  relations  of  proteid  metabolism  ;  American  Journal  of 
Physiology,  1903,  X.,  p.  115. 

7.  A  study  in  the  course  of  the  nitrogen,  sulphate  and  phosphate  excre- 
tion, as  observed  in  short  periods  following  a  small  increase  in  the  proteid 
ingested  (with  Joseph  S.  Chamberlain);  American  Journal  of  Physiology, 
1904,  X.,  p.  269. 

8.  On  the  morphological  changes  in  the  blood  after  muscular  exercise  ; 
American  Journal  of  Physiology,  1904,  X.,  p.  384. 

9.  On  the  influence  of  external  hemorrhage  on  chemical  changes  in  the 
organism,  with  particular  reference  to  proteid  catabolism  (with  W.  J.  Gies); 
American  Journal  of  Physiology,  1904,  XI.,  p.  171. 

10.  On  the  influence  of  copious  water  drinking  ;  University  of  Pennsyl- 
vania Medical  Bulletin,  1905,  XVIII.,  p^  7. 


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59 


TABLE  v.— ELIMINATION  OF  URINE 
As  Influenced  by  Anaesthesia  and  Hemorrhage 


1/  0 

II 

Experiment. 

Date. 

Time 
Begun. 

First  urine 
passed. 

Hour  s 
operatioi 
u  r  i  ti  e 
passed. 

Remarks- 

1902 

a.  m. 

p.  m. 

First    hemor- 
rhage   Inov.  14 

(2.93  percent.) 

8.10 

Between  Mid- 
night and  S.30 
a.  m. 

(Nov.  15th) 

16 — 24 

300 

Anaesthesia . . .  Nov.  30 

9  10 

4- 15 

7 

283 

Anaesthesia  — 
operation  . . .  Dec.  10 

8.12 

4.20 

8 

294 

Second  hemor- 
rhage   

(3.22  per  cent.) 

Third     hemor- 
rhage   

(351  per  cent.) 

Dec.  23 

1903 
Jan.  13 

8.14 
8.20 

Between  Mid- 
night and  9 

a.  m. 
(Dec.  24) 

1-2 

16 — 25 
5-6 

590 
180 

fNo  urine  8  hours  before 
the    operation.       Hence 
1  the  major  portion  of  the 
■  180   cc.    passed   between 
I  p.  m.  and  2.  p.  m.  was 
evidently  in  the  bladder 
at  the  time  of  operation. 

Fourth  hemor- 
rhage   Jan.  20 

(3.26  per  cent.) 

8.29 

8.30 

12 

200 

f  No  more  urine  until  after 
(I  p.  m.  Jan.  2i.st.           - 

Fifth     hemor-  [ 

rhage    i  Jan.  24 

(2.46  per  cent.)i 

8.30 

2.20 

6 

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M*' 

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«  C 

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c 

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a 

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t:  u 

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1 
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2S 

2  u 

Cu 

w 

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66 


TABLE  XII. 

Daily  Data  of  the  First  Series  of  Experiments. 

/.  Preliminary  Experiment,  Nov.  2-1  j,  igo2 


Body- 
weight. 

Urine. 

Feces. 

Day 

No. 

Volume. 

Specific 
gravity. 

Nitrogen. 

Period  average  to  date. 

Weight. 

Volume. 

Nitrogen. 

Fresh. 

Dry. 

kilos 

cc. 

grams 

cc. 

grams 

grams 

grams 

I 

16.96 

499 

1017 

8.30 

499 

8.30 

46.5 

24.4 

2 

16.96 

581 

1019 

10.23 

540 

9.26 

3 

16.96 

394 

I02I 

7.82 

491 

8.78 

43-1 

22.1 

4 

16.95 

499 

1020 

9.84 

493 

9-05 

.^5-8 

19.7 

5 

16.92 

461 

1020 

8.56 

487 

8.95 

27.8 

15-6 

6 

16.89 

445 

1020 

8.10 

480 

8.81 

71. 1 

36.4 

7 

17.01 

400 

IO18 

6.51 

468 

8.48 

8 

16.78 

662 

IOI8 

11.27 

493 

8.83 

54.1 

.38.1 

9 

16.91 

436 

IOI8 

7-33 

486 

8.66 

32.4 

18.2 

10 

16.83 

443 

IOI9 

7.71 

482 

8.57 

30.6 

17.7 

II 

16.85 

486 

IOI9 

8.57 

482 

8.57 

32.3 

17.8 

12 

16.80 

560 

IOI9 

10.60 

489 

8.74 

27.2 

14.4 

//.  First  Hemorrhage,  (2.93  per  cent.)  Nov.  f4-2g,  igo2 


13 

16.29 

377 

1017 

6.33 

377 

6.33 

34-3 

20.2 

14 

16.20 

556 

1027 

13-34 

467 

9.84 

15 

]6.20 

501 

1022 

10.87 

478 

10.18 

36-1 

20.5 

16 

16.25 

370 

1022 

763 

451 

9-54 

69.1 

34-1 

17 

16.20 

570 

1025 

13-67 

475 

10.37 

18 

16.32 

•  307 

1017 

5-25 

447 

9-51 

45-3 

23-1 

19 

16.37 

475 

1022 

9.84 

451 

9-56 

33-6 

20.0 

20 

16.33 

586 

1021 

11.40 

468 

9-79 

38.2 

19.0 

21 

16.34 

462 

1020 

9.69 

467 

9.78 

41.8 

27-5 

22 

16.33 

512 

1019 

8.84 

472 

9.69 

23 

16.32 

520 

1018 

8.20 

476 

9-55 

37-5 

19-5 

24 

16.28 

498 

1019 

9.41 

478 

9-54 

37-1 

20.7 

25 

16.29 

463 

102 1 

9.28 

477 

9-52 

35-8 

17.4 

26 

16.27 

444 

1019 

7-56 

474 

9-38 

42.5 

21.6 

27 

16.18 

576 

1019 

IO-33 

481 

9-44 

40.3 

20.0 

28 

16.09 

615 

1020 

11.04 

489 

9-54 

32-8 

16.9 

^ 

///.  Anaesthesia,  Nov. 

SO- Dec.  9 

igo2 

29 

i5-8i 

654 

1018 

8.59 

654 

8.59 

64.0 

35-9 

30 

15-85 

495 

1021 

8.56 

575 

8.58 

31 

15.88 

467 

1019 

8.80 

539 

8.65 

36.2 

20.1 

32 

16.02 

388 

1021 

8.22 

501 

8.54 

31.3 

16.7 

33 

16.03 

512 

1020 

10.70 

503 

8.97 

59-3 

32.6 

34 

16.09 

458 

loiq 

8.06 

496 

S.82 

35 

16.00 

570 

1018 

9.01 

506 

8.85 

64.0 

36.4 

36 

16.00 

555 

1018 

8.78 

512 

8.84 

32.6 

17-3 

37 

15-98 

593 

1017 

9.20 

521 

8.88 

38 

15-94 

532 

1020 

9-65 

522 

8.96 

71.9 

35-3 

^7 


TABLE  XII     {Co7iti7iued) 
/K  Anaesthesia^Operation,  Dec.  10-22,  igo2 


Body- 
weight. 

Urine. 

Feces. 

Day 

No. 

Vohime. 

Specific 
gravity. 

Nitrogen. 

Period  average  to  date. 

Weight. 

Volume. 

Nitrogen. 

Fresh. 

Dry. 

kilos 

cc. 

grams 

cc. 

grams 

grams 

grams 

39 

15-44 

754 

I018 

9-98 

754 

9-98 

40.0 

20.5 

40 

15-42 

572 

1020 

10.72 

663 

10.35 

41 

15.64 

330 

I019 

6-73 

552 

9-15 

35-2 

19.0 

42 

15-67 

520 

1024 

12.28 

544 

9-93 

28.1 

15-5 

43 

15-68 

530 

1020 

10.07 

541 

9.96 

63-4 

35-4 

44 

15-68 

596 

IOI8 

9.27 

550 

9.84 

45 

15-72 

400 

IOI9 

7-77 

529 

9-55 

41.9 

21.7 

46 

15-69 

558 

1020 

II. 12 

533 

9-75 

I18.8 

51-4 

47 

15-68 

526 

1023 

12.81 

532 

10.09 

48 

15-66 

556 

I02I 

10.87 

534 

10.16 

21-3 

15-7 

49 

15-74 

405 

IO18 

7-41 

522 

9.91 

32.5 

15-7 

50 

15-70 

575 

IOI8 

9.68 

527 

9.89 

51-0 

29-3 

51 

15-72 

5S0 

IO18 

10.00 

53r 

9.90 

23.2 

13-3 

l^.  Second  Hemorrhage  {j. 22  per  cent.)  Dec.  2j,  igo2-Jan.  12,  /goj 


52 

15-18 

428 

1014 

5.58 

428 

5-58 

30.3 

19.1 

53 

15-09 

590 

1027 

13-51 

509 

9-55 

54 

14.92 

579 

1024 

13.06 

532 

10.72 

30.6 

16.8 

55 

14.76 

620 

1020 

11.62 

554 

10.94 

57-8 

37-7 

56 

14.76 

478 

1026 

II. 89 

539 

II. 13 

26.5 

18.3 

57 

14.76 

447 

1024 

10.27 

524 

10.99 

18.4 

II. I 

5« 

14-77 

534 

1024 

13-19 

525 

11.30 

27.8 

14-7 

59 

14.77 

482 

1022 

10.93 

520 

ir.26 

41-5 

20.7 

60 

14.78 

415 

lorS 

7.02 

508 

10.78 

48.7 

24-5 

61 

14-75 

566 

1023 

11.77 

514 

10.88 

24.3 

16.7 

62 

14.70 

598 

1019 

10.29 

52r 

10.83 

33-5 

16.9 

63 

14.64 

600 

1020 

11.47 

528 

10.88 

31.8 

16.8 

64 

14.56 

534 

1020 

10.33 

529 

10.84 

69.0 

37-7 

65 

14.44 

620 

1019 

10.97 

535 

10.85 

66 

14-46 

468 

1020 

9-48 

531 

10.76 

35-5 

17-5 

67 

14-45 

546 

1021 

10.97 

532 

10.77 

69.1 

35-9 

68 

14-43 

522 

1019 

9.61 

5.^t 

10.70 

69 

14-45 

510 

1020 

9-84 

530 

10.65 

48.9 

26.3 

70 

14-38 

542 

1019 

972 

531 

10.61 

33-5 

21.0 

71 

14.40 

518 

io;8 

9  22 

530 

10.54 

37-8 

19.8 

72 

14.42 

534 

1018 

9.17 

530 

10.47 

103.8 

46.1 

68 


85 


TABLE  XII     {^Continued^ 
VI.   Third  Hemorrhage  {3.51  per  cent. )  Jan.  13-19,  1903 


Body- 
weight. 

Urine. 

Feces. 

Day 

No. 

Volume. 

Specific 
gravity. 

Nitrogen. 

Period  average  to  date. 

Weight. 

Volume. 

Nitrogen. 

Fresh. 

Dry. 

kilos 

cc. 

grams 

cc. 

grams 

grams 

grams 

73 

13.76 

460 

IOI8 

7.91 

460 

7.91 

74 

13.80 

475 

1026 

11-55 

468 

9-73 

... 

75 

13-70 

624 

I02I 

13.01 

520 

10.82 

47-5 

23.0 

76 

13.76 

394 

1023 

9-03 

488 

10.38 

42.3 

21.7 

77 

13.82 

412 

1024 

10.44 

473 

10.39 

57-8 

28.0 

78 

13.80 

484 

1024 

10.96 

475 

10.48 

103.9 

38-5 

79 

13-77 

488 

IOI8 

8-33 

477 

10.18 

56.5 

19.7 

VII.  Fourth  Hemorrhage  {3.26 per  cent.)  Jan.  20-23,  1903 


80 

13-35 

463 

IOI8 

8.88 

463 

8.88 

81 

13.36 

518 

1033 

13-92 

490 

11.40 

82 

13-04 

770 

1022 

17-14 

584 

13-31 

29-5 

15-9 

83 

12.90 

464 

1024 

11-75 

554 

12.92 

134-5 

59-6 

VIII.  Fifth  Hemorrhage  {2.46 per  cent.)  Jan.  24-23,  1903 


11.28     I      ..-.      I      ....      I  ..••    I   •■•• 

Death  occurred  at  the  end  of  the  .<^econd  hour. 

I  I  II 


69 


TABLE  XIII 
Relation  Between  Total  Nitrogen  and  Volume  of  Urine. 


Preliminary  experiment-  ■  •  • 

After  first  hemorrhage 

(2.93  per  cent. ) 

Preliminary  to  Anaesthesia 
Experiment 

(Dog  in  nitrogen  equil. ) 

First  four  days  of  Anaesthe- 
sia experiment 

(Control  experim't  No.  i ) 

Last  four  days  of  Anaesthe- 
sia experiment 

(Dog  in    nitrogen  equili- 
brium) 

1  irst  five  days  of  Anaesthe- 
sia-operation experiment- 
(Control  experim't  No.  2) 

Preliminary  to  .second  hem- 
orrhage   

(Dog  in  nitrogen  equil.) 

After  second  hemorrhage-  -  • 
(3.22  per  cent. ) 

Preliminary  to  third  hem- 
orrhage   

(Dog  in  nitrogen  equil.) 

After  third  hemorrhage  •  ■  -  • 
(3.51  per  cent.) 

After  fourth  hemorrhage  - .  ■ 
(3.26  per  cent. ) 

After  fifth  hemorrhage 

(  2.46  per  cent.) 


Nov.  2-13 
1902 

Nov.  14-18 


Nov.  25-29 

Nov.  30-Dec.  3 

Dec.  6-9 

Dec.  10-14 

Dec.  19-22 
Dec.  23-29 

Jan.  S-12 
1903 

Jan.  13-15 

Jan.  20-21 

Jan.  24 


No.  of 
days. 


Average 
Nitrogen 
(grams) 

Average 

volume. 

cc. 

8.74 

489 

10.37 

475 

9-52 

519 

8.54 

501 

9.16 

563 

9.96 

541 

9-49 

529 

11.30 

525 

9-51 

525 

10.82 

520 

11.40 

490 

11.28 

455 

Ratio. 
(N:V) 


1:55-9 
1:45-8 

1:54-5 

1:58.7 
1:61.5 

1:54-3 

1:55-7 
1:46.5 

1:55-2 

1 :48. 1 
1:43.0 
1:40.3 


JO 


TABLE  XIV 

Nitrogen  Content  of  Composite  Urine  Samples. 


Experiment. 

No.  of 
days. 

Nitrogen  (grams). 

tn-r; 

Si  s 

Content  of 

composite 

urine  sample. 

Total  content 
of  daily 
samples. 

Daily 
variation. 

12 
16 
ID 

13 
21 

7 
4 

104.30 

^52-31 
89.48 

130.32 
220.48 

70.54 
51. r6 

104.84 
152.68 

89-57 
128.70 
219.91 

71-23 
51-69 

0.045 
0.023 
0.009 
0.125 
0.027 
0.098 
0.142 

First  hemorrhage 

CO 

.M 

a; 
cn 

s 

Anaesthesia-operation 
Second  hemorrhage  •  • 

Third  hemorrhage 

Fourth  hemorrhage- . . 

'j*otal 

83 

818.59 

818.62 

003^ 

tn 

9 
9 

5 

69.92 
78.82 
46.82 

69-57 
79.11 

47-34 

0.039 
0.032 
0.104 

■73 
O 

Anaesthesia-operation 
Hemorrhage 

Total 23 

195-56 

196.02 

0.46 

1  This  variation  of  but  0.03  gram  of  nitrogen  for  a  period  of  83  days  is  very  striking. 


7T 


TABLE  XV 

Daily  Diet  (Second  Series). 


Constituent  of 
the  diet. 


Beef 

Cracker  dust 

Lard 

Bone  ash  . .  • 
Water 

Total 


<  be 


200 

52 
15 

8 
375 


650 


Nitrogen. 


Per 
cent. 


3-759 
1-55 
0.028 
0.026 


Grams. 


7-5'8 
0.806 
0.004 
0.002 


8.33 


Sulphur. 


Per 
cent. 


0.288 
O.1318 
0.03 
0.06 


0.576 
0.069 
0.004 
0.005 

0.654 


Phosphorus. 


Per 
cent. 


0.220 

0.1345 
0.085 

17-79 


Grains. 

0.440 
0.070 
0.013 
1-423 


1.946 


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o 

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c 
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to    ^ 
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m 


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T 

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00 

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00 

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d 

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CJN 

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fc 

1 

1 

1 

1 

i 

1 

0 

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r^           0) 

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Pi 

V 

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m               fo 

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8 

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oc 

00 

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cc 

CO 

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1 

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jj 

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b. 

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0; 

>> 

ca  a 

ca  P- 

11 

s 

0; 

0  .  s 

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_s 

ca 

s  o 
ca  1 

S  0 
ca  1 

s 

0 

S:" 

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ca 

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C3  , 

V 

•"^  <u  u 

OJ 

p 

w  iJ  o      u 

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■<-.53  ca     i*- 

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OJ  t- 

Ps 

pq 

< 

< 

P5 

73 


TABLE  XVII 

Dailv  Data  of  the  Second  Series  of  Experiments. 

/.  Ptelifninary  Experiment,  June  2-10,  igoj 


Body- 
weight. 

Urine. 

Feces. 

Day 
No. 

Volume. 

Specific 
gravity. 

Nitrogen. 

Period  average  to  date. 

Weight. 

Volume. 

Nitrogen. 

Fresh. 

Dry. 

kilos 

cc. 

gram.s 

cc. 

gram.s 

grams 

grams 

I      1 

1 1. 85 

390 

I016 

7.63 

390 

7-63 

.22.8 

15.8 

^      i 

11.80 

415 

IO18 

6.99 

40,^ 

7 

31 

... 

3     1 

11.82 

388 

1020 

7.58 

398 

7 

40 

50.2 

293 

4 

11.70 

485 

1020 

965 

419 

7 

96 

5 

11.76 

344 

I017 

5-31 

404 

7 

43 

6 

11.60 

515 

I018 

963 

423 

7 

80 

40.3 

30.8 

7 

11-57 

402 

IOI9 

8.14 

420 

7 

85 

34.2 

21.4 

8 

11.63 

298 

IOI8 

5-52 

405 

7 

56 

40.0 

235 

9 

11.68 

452 

1020 

9.12 

410 

7 

73 

41.5 

24-3 

//.  Anaesthesia — operation,  June  ii-ig,  /goj 


10 

11.40 

450 

1019 

8.56 

450 

8.56 

54-8 

31.8 

II 

11.30 

560 

1019 

9.46 

505 

9.01 

12 

11.28 

484 

1018 

9-33 

498 

9.12 

22 

.5 

17.9 

13 

11.24 

432 

1019 

8.66 

482 

9.00 

14 

11.26 

422 

1016 

6.3t 

470 

8.46 

57 

5 

28.5 

15 

II. 10 

544 

1023 

•2.55 

482 

9.14 

16 

11.06 

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,,,§PUTH  PROPERTY,^, 

Hawk 

The  influence  of  hemorrhage  upon 
metabolism 


MAY  12,1943      J^c^^-Slj^iJ^ 


DM  PERSONAL  RLSh:KVii:  SHELF 


